EDINBURGH JOURNAL OF BOTANY 63 (2&3): 161–182 (2006) 161 doi:10.1017/S0960428606000631 E Trustees of the Royal Botanic Garden Edinburgh (2006) Issued 30 November 2006 A SYNOPSIS OF THE GENUS BERLINIA (LEGUMINOSAE – CAESALPINIOIDEAE) B. A. MACKINDER1 & D. J. HARRIS2 BerliniaSol.exHook.f.(Leguminosae–Caesalpinioideae)isunderinvestigationwithtwo maingoals:(i)todelimitanddescribethespecies,and(ii)toelucidatetheirrelationships. Delimitationanddescriptionisnowlargelycompletedandispresentedinthissynopsis. ThegenusBerliniaisassessedascomprising18species.Keystothespeciesarepresented andfor eachspeciesa brief description, selected synonymy and specimencitations are given. Amonograph ofBerlinia including afull taxonomictreatment andthe results of thephylogenetic investigation will bepublished later bythe seniorauthor. Keywords.Berlinia,Caesalpinioideae,Fabaceae,Leguminosae,taxonomy,tropicalAfrica. INTRODUCTION BerliniaisconfinedtotropicalAfricaandwasdescribedin1849byJ.D.Hookerwho created the genus to accommodate a single species. Establishing authorship of the genuswasnotstraightforward.IntheNigerFlora,theentryforBerliniasimplyreads BerliniaSoland.ThemanuscriptnameBerliniaacuminataiswrittenononeofthethree differentcollections(Smeathmanns.n.)onwhichthegenusisbasedinwhatisthought to be Solander’s handwriting (Keay, 1954). Although the Niger Flora was edited by W. J. Hooker, in the preface, he credits the majority of the work to his son J. D. Hooker.However,atthebeginningoftheaccountoftheLeguminosae,afootnotesays ‘by J. D. Hooker and G. Bentham’. Nevertheless, throughout the Leguminosae accountauthorshipsofnewgeneraareattributedtooneorotherauthor,e.g.Hook.f. to Ostyrocarpus, or Benth. to Leucomphalus. As the primary literature is somewhat ambiguouswehavebeenobligedtoassessevidencefromsecondarysources.Inalater paperBentham(1866)describedtwoadditionalnewspeciesinthegenus(B.auriculata Benth.andB.bracteosaBenth.),buttooknocreditfortheauthorshipofB.acuminata, simplycitingthespeciesasB.acuminataSoland.,inHook.f.Subsequently,whenKeay (1954) unravelled the taxonomic and concomitant nomenclatural complexities of B. acuminata,hestatedthatBerliniawasdescribedbyJ.D.Hooker.Wethereforebelieve thatthegenus wasdescribedby J.D. Hooker,usingSolander’smanuscriptname so thatthegenusiscorrectlyattributedtoSol.exHook.f. By the early 1900s, 26 more species had been described and the generic limits of Berlinia s.l. had become unclear. During the first half of the 20th century, several authors (Harms, 1915; De Wildemann, 1925; Baker, 1930; Pellegrin, 1949; 1RoyalBotanicGardens,Kew,Richmond,SurreyTW93AB,UK.E-mail:[email protected] 2RoyalBotanicGardenEdinburgh,20AInverleithRow,EdinburghEH35LR,UK. 162 B. A. MACKINDER & D. J. HARRIS Duvigneaud,1950;Troupin,1950;Hauman,1952a)proposedavarietyoftaxonomic systems, to accommodate the considerable morphological diversity of Berlinia s.l., sometimes referred to as the Berlinia complex. Eleven new segregate genera were described. Despite the wealth of literature, there was no consensus on the generic limits of Berlinia and in consequence taxonomic confusion surrounded Berlinia and its putatively allied generic segregates. In 1957, Le´onard published a higher level tribal account in which he proposed a narrower circumscription for Berlinia. Le´onard’s delimitation was based on Harms’ (1915) sections Euberlinia and Macroberlinia and gained wide acceptance by the authors of Berlinia accounts in the major African Floras, namely Torre & Hillcoat (1956) in Conspectus Flora Angolensis, Keay (1958) in Flora of West Tropical Africa, Brenan (1967) in Flora of EastTropicalAfrica,Aubre´ville(1968and1970,respectively)inFloreduGabonand FloreduCameroun.Hauman(1952b)raisedsectionMacroberliniatogenericrankin Flore du Congo Belge et du Ruanda-Urundi. There is significant disagreement between some of these floristic accounts of the genus which we believe can be resolved by revising Berlinia over its entire distribution. Thispaperpresentstheprogressthathasbeenmadetowardsclarifyingthespecies- level taxonomy of the genus. Eighteen species are recognized, including two species recently described (Mackinder, 2006), of which 16 are assigned to section Berlinia. Within this section, species are seldom distinguished by a unique character or characters but instead species delimitation frequently rests on unique combinations of characters. Our findings confirm Brenan’s (1963) observation that the species of this section ‘are all uncommonly closely related to one another’. The primary aim of this synopsis is to provide a means to identification. A brief descriptionofthegenus,thecharactersdifferentiatingthetwosections(Table1)and keystothespeciesbasedonherbariummaterialandpodsarepresented.Toassistin the use of the former key, the comparative states of four characters have been illustrated (Fig. 1). Geography is included in the key where it can assist the user in choosing the correct option. Pending the results of a phylogenetic investigation, the speciesaretreatedinalphabeticalorderwithinthetwosections.Foreachspeciesthe accepted name with a brief synonymy is given, followed by geographical distribu- tion, citation of selected specimens and a short description of recorded habitats. SYNOPSIS OF BERLINIA NOM. CONSERV. Berlinia Sol. ex Hook.f. in Hook., Niger Flora: 326–328 (1849), nom. conserv. – Type: Berlinia grandiflora (J.Vahl) Hutch. & Dalziel. Westia J.Vahl, Skr. Naturhist.-Selsk. 6: 117 (1810). – Type: Westia grandiflora J.Vahl. Small to large forest trees, rarely shrubs. Leaves compound in (1–)2–5(–6) pairs, lower pair often shorter and broader. Inflorescence axillary or terminal, a solitary robust (or rarely slender) erect raceme, a cluster of racemes or a few to A SYNOPSIS OF THE GENUS BERLINIA 163 TABLE 1.Characters thatdistinguish Berliniasections Berlinia and Macroberlinia Berlinia section Berlinia BerliniasectionMacroberliniaHarms 1. Inflorescence commonly a fewto Inflorescence a solitary many-branched panicle, sometimes a clusterof robusterect raceme racemesorveryrarely a slender, solitaryraceme 2. Bracts inconspicuous,early caducous, Bracts conspicuous, semi-persistent seldomseen to persistent,often seen 3. Petaldevelopment unequal; adaxial petal Petaldevelopment subequal well developed, laterals andabaxials poorly developed 4. Maturepods commonly denselygolden Mature podsglabrous,drying puberulous,more rarelyglabrous anddrying black andslightlyshiny dull orshiny many-branched panicle. Bracts conspicuous and semi-persistent to persistent or inconspicuous,soonfalling.Bracteoleslarge,fullyenclosingthebudbeforeanthesis. Hypanthium conspicuous, tubular, 9–22 mm long. Sepals 5, reflexed. Petals 5; petal development subequal, or adaxial petal well developed but laterals and abaxials poorly developed. Stamens 10, 9 fused at the base into a ring, 1 free. Staminodes absent.Podsupto50cmlong,woody,laterallycompressed,2-valved,dryinggolden to dark brown or black, glabrous or hairy, explosively dehiscent, terminal suture broad, up to 18 mm wide. Seeds 2–7. Seedling germination epigeal. Ofthe18species,17arefoundintheGuineo-Congolianregion,withasoleoutlierin the coastal forests of southern Tanzania and northern Mozambique. Key to the sections 1a. Petalsofsimilarlength,thelaminaoftheadaxialpetalbroadest;bractslarge,at least 22 mm long, conspicuous in the maturing inflorescence, sometimes persistent until the flowers have opened; immature and mature fruits glabrous, drying black and slightly shiny _____ section Macroberlinia (species 1 and 2) 1b. Petals very unequal in length, the adaxial petal broader and longer than the others, the lateral petals barely as long as the claw of the adaxial petal; bracts small,notexceeding5mmlong,fallingwhentheinflorescenceisveryimmature; immatureandmaturefruitscommonlydenselygoldenpuberulous,occasionally glabrous and then drying golden and shiny or dull black or brown ________ _________________________________________ section Berlinia (species 3–18) Key to the species of section Macroberlinia 1a. Bracts 45–105 mm long, enclosing and obscuring the unopened part of the inflorescence to give a crowded ‘head-like’ appearance, then falling when the 164 B. A. MACKINDER & D. J. HARRIS A SYNOPSIS OF THE GENUS BERLINIA 165 flowers open; bracteoles 52–75 mm long, glabrous or sparsely puberulous on theinterior surface,thesurfacenotconcealed bytheindumentum;widthofthe adaxial (widest) petal 55–92 mm; known from Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola _______ _______________________________________________________1. B. bracteosa 1b. Bracts 22–30 mm long, enclosing only very immature buds then falling, more maturebudsclearlyvisibleintheunopenedpartoftheinflorescence;bracteoles 22–38 mm long, densely puberulous on the interior surface, the surface concealed by the indumentum; width of the adaxial (widest) petal 55–68 mm; known from Sierra Leone, Liberia, Ivory Coast and Ghana ______________ _____________________________________________________ 2.B.occidentalis Key to the species of section Berlinia 1a. Hypanthium glabrous, or sparsely and unevenly white pubescent, or the white pubescence present in one or more irregular narrow longitudinal bands, sometimes the band becoming broader and meeting towards the base of the hypanthium,most or all ofthesurface ofthe hypanthiumnot concealedby the indumentum, the indumentum not extending to the sepals _______________ 2 1b. Hypanthium golden tomentose, either completely covered or the indumentum confined to one broad longitudinal band, the surface of the hypanthium concealed by the indumentum (where present), the indumentum sometimes extending to the sepals ____________________________________________ 12 2a. Ovary moderately villous, the surface not concealed beneath; known only from Gabon _______________________________________________ 15. B. rabiensis 2b. Ovary densely sericeous or densely puberulous, the surface concealed beneath; widespread ________________________________________________________ 3 3a. Ovary densely sericeous, the individual hairs clearly visible at 6 10 magnification _____________________________________________________ 4 FIG. 1. Berlinia section Berlinia floral ground plan with comparative states of four morphologicalcharactersinBerlinia.A,flower;B,ovarysurfacedenselypuberulous;C,ovary surface densely sericeous; D, ovary surface moderately villous; E, hypanthium surface with whitish pubescence in irregular narrow longitudinal bands; F, hypanthium surface partially concealedbyasinglebroadgoldentomentoseband;G,hypanthiumsurfacefullyconcealedby a golden tomentose indumentum; H, bracteole interior surface concealed by densely silky indumentum; J, bracteole interior surface concealed by densely villous indumentum; K, bracteoleinteriorsurfaceconcealedbydenselypuberulousindumentum;L,bracteoleinterior surface only partially concealed by moderately puberulous indumentum; M, bracteole thick; N,bracteolethin.AfromLisowski132,BfromLouis2322,CfromLatiloFHI34423,Dfrom Schoenmaker 330, E from Lisowski 132, F from Harley 1494, G from Preuss 189, H from Latilo FHI 34423, J from Gossweiler 13745, K from Louis 2322, L from Touissant 2192, M from Lisowski 132, N from LatiloFHI34423.Drawn byMargaretTebbs. 166 B. A. MACKINDER & D. J. HARRIS 3b. Ovary densely puberulous, the individual hairs not visible at 6 10 magnification _____________________________________________________ 9 4a. Bracteole interior glabrous, sparsely to densely puberulous, the surface concealed or not by the indumentum _________________________________ 5 4b. Bracteole interior densely villous or silky, the surface concealed by the indumentum _____________________________________________________ 11 5a. Bracteoles11–1966–8mm;pedicels7–12mmlong;inflorescenceaxesglabrous to sparsely puberulous; known from Nigeria, Cameroon, Gabon and Congo (Brazzaville) __________________________________________ 3. B. auriculata 5b. Bracteoles 22–51 6 8–20 mm; pedicels 14–60 mm long; inflorescence axes moderately to densely puberulous ____________________________________ 6 6a. Hypanthium indumentum present, usually arranged in irregular longitudinal bands of pubescence, sometimes with a more complete covering but always somepartofthesurfaceclearlyvisible;bracteoles45–51617–22mm;pedicels 24–60 mm long, leaflets concolorous, the tertiary venation visible and raised; known from Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola (Cabinda) __________________________________________ 18. B. viridicans 6b. Hypanthiumglabrousorwithscatteredandsparsewhitepubescence;bracteoles 22–5068–15mm;pedicels14–36mmlong,leafletsconcolorousordiscolorous, the tertiary venation visible and raised or not __________________________ 7 7a. Tertiaryleafletvenationonbothsurfacesvisibleandraised,leaflets(1)2–3pairs, usually concolorous and drying green; known from Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola ______________________________________________ 6. B. congolensis 7b. Tertiary leaflet venation on both surfaces indistinct, leaflets (2)3–5 pairs, discolorous or concolorous, usually drying brown ______________________ 8 8a. Stylenotexceeding30mmlong;undersideofleafletsglabrous;adaxialpetal35– 48 mm long, bracteoles 8–13 mm wide, the interior surface moderately to densely puberulous, the surface only partially concealed by the indumentum; tree of coastal forest on sandy soil; known from Tanzania and Mozambique _ ______________________________________________________ 13.B.orientalis 8b. Style more than 40 mm long; underside of leaflets glabrous; adaxial petal 44– 62mmlong,bracteoles10–15mmwide,theinteriorsurfacedenselypuberulous, the surface visible in places; tree of coastal forest, known from Sierra Leone, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea and Gabon ___ ________________________________________________________ 5.B.confusa 9a. Bracteoles 2–4 mm thick (see Note 1); 15–32 mm wide _________________ 10 9b. Bracteoles thinner, c.1 mm thick; 9–18 mm wide; known from Congo (Brazzaville) and Congo (Kinshasa) _____________________ 14. B. phenacoa A SYNOPSIS OF THE GENUS BERLINIA 167 10a. Leaflet length 3 or more times the width, leaflets often slightly falcate, upper pair of leaflets opposite, lower pairs opposite or sub-opposite; hypanthium unevenly pubescent, the surface clearly visible between irregular longitudinal bandsofhairs;pedicels23–60mmlong;petiolules5–18mmwide;knownfrom Cameroon, Equatorial Guinea, Central African Republic, Gabon, Congo (Brazzaville) and Congo (Kinshasa) _______________________ 4. B. bruneelii 10b. Leaflet length not more than 2K times the width, leaflets straight, all pairs of leafletsopposite,hypanthiumglabrous,occasionallywithpatchesofpubescence atapexorwithsparsescatteredpubescence;pedicels18–25mmlong;petiolules 7–12mmlong;knownfromNigeriaandCameroon __________ 7.B.coriacea 11a. Bracteole interior densely silky, indumentum pale, soft to the touch and often with a slight sheen; petiolules 5–9 mm; upper leaflets 7.0–22.0 6 3.2–11.5 cm; petiole 20–42(–65) mm; bracteole width 11–20 mm; known from Mali, Guinea (Conakry), Niger, Sierra Leone, Ivory Coast, Ghana, Togo, Benin, Nigeria, Equatorial Guinea, Cameroon, Central African Republic and Congo (Kinshasa) _________________________________________ 10. B. grandiflora 11b. Bracteole interior densely villous, indumentum mid brown, not soft to the touch,withoutasheen;petiolules3–4mm;upperleaflets7.5–9.062.4–4.0cm; petiole 12–20 mm; bracteole width 7–13 mm, known only from Congo (Kinshasa) _____________________________________________ 16. B. sapinii 12a. Adaxial petal length 31–40 mm, adaxial petal width 21–40 mm _________ 13 12b. Adaxial petal length 52–85 mm, adaxial petal width 46–94 mm _________ 14 13a. Pedicels 10–12 mm long; adaxial petal width 21–28 mm; underside of leaflets frequently appearing glabrous or occasionally sparsely puberulous at 6 10 magnification, at higher magnifications appearing sparsely to moderately puberulous; tertiary venation indistinct or visible above and below; leaf apex acuminate, acumen 4–13 mm; bracteoles 18–28 6 8–14 mm, interior surface denselypaletogoldenpuberulous,becomingpatchyonolderbracteoles;known fromSierraLeone,Liberia,IvoryCoastandGhana _______ 17.B.tomentella 13b. Pedicels 30–36 mm long; adaxial petal width c.40 mm; underside of leaflets moderatelypubescentonmid-vein,secondaryveinsandverylowerpartofleaf blade and at higher magnifications; tertiary venation visible and somewhat raised;leaf apex acute; bracteoles 24–40 6 12–20 mm, interior surface densely villous; known from Congo (Kinshasa) and Angola ________ 12. B. lundensis 14a. Bracteolesnotexceeding1mmthick;narrowlyoblong,14–16mmwide;known only from Nigeria _____________________________________ 11. B. hollandii 14b. Bracteoles 2–4 mm thick; obovate, 16–35 mm wide ___________________ 15 15a. Inflorescence a branched panicle; leaf apex acute or short (3 mm) acuminate; adaxialpetal50–75643–72mmwide;knownfromaltitudesof(450–)900–1700m; knownfromCongo(Kinshasa),Burundi,AngolaandZambia _______________ ___________________________________________________________ 9.B.giorgii 168 B. A. MACKINDER & D. J. HARRIS 15b. Inflorescence acluster of2–4 racemes,rarelyasolitary raceme; leaf apexshort to long acuminate, acumen (5–)7–12 mm; adaxial petal 67–85 6 55–94 mm wide; known from altitudes of 1–400 m; known from Nigeria, Cameroon and Gabon _______________________________________________ 8. B. craibiana Key to mature Berlinia pods The majority of Berlinia species cannot be distinguished in fruit alone. However, a third of the species, including some common ones, can be recognized from fallen valvesandsoapodkeyisincludedhere.ThepodsofBerliniahollandiifromNigeria are unknown. 1a. Pods covered in a dense felt of pale gold, gold to reddish-gold indumentum 4. B. bruneelii; known from Cameroon, Equatorial Guinea, Central African Republic, Gabon, Congo (Brazzaville) and Congo (Kinshasa). 6. B.congolensis;knownfromNigeria,Cameroon,EquatorialGuinea,Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola. 8. B. craibiana; known from Nigeria, Cameroon and Gabon. 9. B. giorgii; known from Congo (Kinshasa), Burundi, Angola and Zambia. 10. B. grandiflora; Mali, Guinea (Conakry), Niger, Sierra Leone, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Equatorial Guinea, Central African Republic and Congo (Kinshasa). 12. B. lundensis; known from Congo (Kinshasa) and Angola. 13. B. orientalis; known from Tanzania and Mozambique. 14. B. phenacoa; known from Congo (Brazzaville) and Congo (Kinshasa). 16. B. sapinii; known only from Congo (Kinshasa). 17. B. tomentella; known from Sierra Leone, Liberia, Ivory Coast and Ghana. 18. B. viridicans; known from Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola (Cabinda). 1b. Podsglabrousorappearingsotothenakedeyeandat610magnification,the surfaces drying golden brown, mid-brown, dark brown or black __________ 2 2a. Pods drying dark brown to black ____________________________________ 3 2b. Pods drying golden brown or mid-brown _____________________________ 5 3a. Pods glabrous at all magnifications, surface appearing dull or slightly shiny _ __________________________________________________________________ 4 3b. Pod appearing glabrous at 6 10 magnification but at higher magnification minutelyreddishpuberulous,surfaceisdull;knownfromSierraLeone,Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea and Gabon ___ ________________________________________________________ 5.B.confusa 4a. Pod surface glossy ________ 1. B. bracteosa or 2. B. occidentalis (see Note 2) 4b. Pod surface dull; known only from Gabon ________________ 15. B. rabiensis A SYNOPSIS OF THE GENUS BERLINIA 169 5a. Pod surface glossy with clearly visible transverse veins; known from Nigeria, Cameroon, Gabon and Congo (Brazzaville) _______________ 3. B. auriculata 5b. Podsurfacedullandscurfywithoutclearlyvisibletransverseveins;knownfrom Nigeria and Cameroon __________________________________ 7. B. coriacea Section Macroberlinia 1. Berlinia bracteosa Benth., Trans. Linn. Soc. London 25: 309 (1866). – Westia bracteosa (Benth.) J.F.Macbr., Contrib. Gray Herb. n.s. 3, 59: 20 (1919). – Macroberlinia bracteosa (Benth.) Hauman, Bull. Se´ances Inst. Roy. Colon. Belge Bull. 23: 518 (1952). – Type: Equatorial Guinea, Bioco, Mann 1434 (holo K!; iso P!). Shrub or tree (2–)8–35 m tall; diameter at breast height (dbh) 11–90 cm. Leaflets in (3–)4–5(–6) pairs; terminal pair (see Note 3) 11.8–44.0 6 4.2–12.6 cm, concolorous or almost so; underside appears glabrous at 6 10 magnification but puberulous at higher magnifications; tertiary venation indistinct above, visible below; apex rounded to shortly acuminate. Petioles 18–64 mm long. Petiolules 8–13 mm long. Inflorescenceasolitaryrobustraceme,axesdenselygoldenbrownpuberulous.Bracts present, 45–105 mm long, persisting in the unopened part of the inflorescence. Bracteoles 52–75 6 15–20 mm, interior surface glabrous or sparsely puberulous. Pedicels 12–20 mm long. Hypanthium glabrous or sparsely and unevenly pubescent. Petals subequal in length; adaxial petal 52–78 6 55–92 mm, apex bilobed. Ovary densely sericeous, individual hairs visible at 6 10 magnification. Immature and mature pod glabrous, drying black and slightly shiny. Distribution and habitat. Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo (Brazzaville), Congo (Kinshasa) and Angola. Lowland forest, forest edges and clearings, along rivers, streams and mangrove edge; 1–650 m. Selection of specimens examined. Nigeria: Calabar, Akampka at the reserve boundary near Awi,fl.19v1965,DaramolaFHI55577(FHI,K!).EquatorialGuinea:Luba–Malabo,km2, fr. 24 iv 1987, Carvalho 2851 (BR!, K!). Cameroon: 16 km on the road from Ebolowa to Minkok, fl. 30 iv 1975, J.J.F.E. de Wilde 8216 (B, BR!, EA, HBG!, K!, LG, MA, MO, P!, PRE, SRGH, WAG!, YA). Gabon: 5–30 km NNW of Ndjole´, fl. 27 iv 1992, Breteler 11138 (G!, WAG!). Congo (Brazzaville): Kouilou, 2˚479S, 11˚519E, fr. undated, Sargos 181 (P!). Congo (Kinshasa): Luki. Perce´e de la Ntosi, fl. 21 v 1952, Gutweiller R.48 (ZT!). Angola: Cabinda, alto Maiombe, fl. 27vi1953, Pinto 355(LISC!). 2. Berlinia occidentalis Keay, Kew Bull. 1954: 269 (1954). – Type: Liberia, Baldwin 11604 (holo K!). Tree(3–)10–35mtall;dbh15–90cm.Leafletsin4–5pairs;terminalpair12.8–25.26 5.2–7.6 cm, concolorous; underside puberulous, hairs just visible at 610 magnification; tertiary venation visible, sometimes raised above and below; apex acute to shortly acuminate. Petioles 15–51 mm long. Petiolules 8–14 mm long. 170 B. A. MACKINDER & D. J. HARRIS Inflorescence a solitary raceme, axes densely reddish brown puberulous. Bracts 22– 30 mm long, present only in very immature inflorescences. Bracteoles 22–38 6 12– 30 mm, interior surface densely puberulous. Pedicels 21–46 mm long. Hypanthium moderately pubescent. Petals subequal in length; adaxial petal 40–70 6 55–68 mm, apexbilobed.Ovarydenselysericeous,individualhairsvisibleat610magnification. Immature and mature pod glabrous, drying black and slightly shiny. Distribution and habitat. Sierra Leone, Liberia, Ivory Coast and Ghana. Fringing forest, commonly along streams and rivers, sometimes in drier areas and on sandy soil; 1–200 m. Selection ofspecimens examined. SierraLeone: Gorahun, fl. & fr.5 vi 1952, Small 725 (BR!, K!,P!).Liberia:NearFirestonesPlantationsalongDukwaiR.,fl.1929,Cooper217(F!,NY!). Ivory Coast: Guiglo, Tai National Park, 2 km ENE of Tai Ecological Station, fr. 5 ix 2001, Wieringa 4322 (WAG!). Ghana: Western Region, close to Aiyinasi, fl. 27 ii 1995, Jongkind 2068(MO, WAG!). Section Berlinia 3. Berlinia auriculata Benth., Trans. Linn. Soc. London 25: 309 (1866). – Type: Cameroon, Mann 2195 (holo K!; iso BM!, GH!, NY!). Shrubortree2–20(–30)mtall;dbh5–77cm.Leafletsin2–3(–4)pairs;terminalpair 7.1–22.4 6 3.5–15.0 cm, discolorous; underside glabrous or occasionally sparsely scatteredpubescent;tertiaryvenationvisible,sometimesraisedabove,visiblebutnot raised below; apex shortly acuminate. Petioles 10–48 mm long. Petiolules 7–10 mm long. Inflorescence commonly a cluster of racemes, rarely a solitary raceme; axes glabrous to sparsely yellowish grey puberulous. Bracts falling early, not seen. Bracteoles thin, 11–19 6 6–8 mm, interior surface glabrous to moderately puberulous, the surface not concealed by the indumentum. Pedicels 7–12 mm long. Hypanthium glabrous. Petals unequal in length; adaxial petal 32–44 6 15–24 mm, apexbilobed.Ovarydenselysericeous,individualhairsvisibleat610magnification. Immature and mature pod glabrous, drying golden or medium brown. Distribution and habitat. Nigeria, Cameroon, Gabon and Congo (Brazzaville). Lowland forest, along rivers and on beaches; 1–500(–900) m. Identificationnote.MaybemisidentifiedasBerliniacongolensis(Bakerf.)Keaywhen sterile but note leaflets of B. auriculata are frequently discolorous (concolorous in B. congolensis). Selection of specimens examined. Nigeria: Degema Distr., fl. ii 1957, Chesters 11 (K!). Cameroon:SecondaryforestnearMundembatown,fl.12v1986,D.W.Thomas6126(BR!,K!, MO, NY!, WAG!). Gabon: Re´serve de la Lope´ au sud d’Ayem, fl. 30 iii 1989, McPherson 13827 (F!, MO, NY!, P!). Congo (Brazzaville): Environs de Brazzaville, foreˆt de la Djoumouna, fl. 27i 1967,Sita 1542(P!, WAG!). 4.Berliniabruneelii(DeWild.)Torre&Hillc.,Bol.Soc.Brot.ser.2,29:40(1955).– BerliniaacuminataSol.exHook.f.var.bruneeliiDeWild.,Ann.Mus.CongoBelge,