A revision of the genus Talbotiella Baker f. (Caesalpinioideae: Leguminosae)

A revision of the genus Talbotiella Baker f. (Caesalpinioideae:Leguminosae)

Barbara A. Mackinder1, Jan J. Wieringa2 & Xander M. van der Burgt1

Summary. A revision of the genus Talbotiella Baker f. (Caesalpinioideae: Leguminosae) is presented. Four speciesfrom Cameroon are described as new and a species of Hymenostegia, also from Cameroon, is transferred to Talbo-tiella bringing the number of species in Talbotiella to eight: T. bakossiensis Cheek, T. batesii Baker f., T. breteleri(Aubrév.) Mackinder & Wieringa, T. ebo Mackinder & Wieringa, T. eketensis Baker f., T. gentii Hutch. & Greenway, T.korupensis Mackinder & Wieringa and T. velutina Burgt & Wieringa. The centre of diversity is Cameroon where sixof the eight species occur, five of which are country endemics. Vegetative characters that can be used to distinguishthe species are provided in tabular form. Species distributions are mapped and all species are assessed for cons-ervation status.

Key Words. Cameroon endemic species, Detarieae, endangered species, Fabaceae, Ghanaian endemic species,Leguminosae, Nigerian endemic species, Tropical Africa.

IntroductionTalbotiella Baker f. (Caesalpinioideae: Leguminosae) isa small genus, principally of forest trees, restricted tothe forests of west and west-central Africa. Prior to thispaper, three species had been described. In 1914,E. G. Baker described Talbotiella eketensis from SENigeria, followed 14 years later by Hutchinson &Greenway who described Talbotiella gentii from Ghana.A year later E. G. Baker added a third species,Talbotiella batesii described only from the type gather-ing made in south-central Cameroon.

However, after examining more recent collectionsmade in Cameroon and Gabon by various collectorsbetween 1972 and 2005, the authors concluded thatthe newer collections represent four undescribedspecies. The correct generic assignment of these newspecies was not immediately clear, so before theycould be described, a detailed evaluation of thegeneric limits of Talbotiella was necessary, particularlywith respect to the neighbouring genus Hymenostegia.Having undertaken such an evaluation based onmorphological, palynological and molecular evidence,it was decided that Talbotiella is a distinct genus andthat the correct placement of all four undescribedspecies is in Talbotiella (Mackinder et al. 2010). Afurther outcome of the generic limits study was thatHymenostegia breteleri Aubrév. was found to be misplacedin Hymenostegia and is here transferred to Talbotiella.

Below, we present a synopsis of Talbotiella enum-erated as comprising eight species, of which one is anew combination and four are described as new. Alleight species in the genus qualify for a category ofthreat as defined by IUCN (2001). Talbotiella ischaracterised by the unique combination of thepresence of imbricate bud scales, persistent petaloidbracteoles, the absence of petals and a pubescentovary. In a few flowers one or two rudimentary linearpetaloid structures were seen but they are nothomologous with the consistently present, two orthree larger spathulate petals of true Hymenostegia.The species of Talbotiella are presented in alphabeticalorder. A map of the distribution of Talbotiella species isshown in Map 1. Species of Talbotiella may bedistinguished from each other vegetatively using thecharacters presented in Table 1. All specimens citedhave been seen by the authors, but not all duplicatescited may have been seen.

TaxonomyTalbotiella Baker f. (1914: 2); J. Léonard (1951: 445),(1957: 125); Cowan & Polhill (1982: 128); Mackinder(2005: 86). Type species: T. eketensis Baker f.

Trees or rarely shrubs (T. eketensis), 6 – 35 m tall, dbh10 – 76 cm. Stipules (seen in young foliage), in pairs,

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010

Accepted for publication September 2010.1 HLAA, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK.2 National Herbarium of the Netherlands, Wageningen University Branch, Biosystematics Group, Generaal Foulkesweg 37, 6703 BL, Wageningen,

The Netherlands.

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free, auriculate at base, the auricle suborbicular, oftenlobed, upper part of stipule narrowly lanceolate orlinear. Leaves alternate, paripinnate, petiole short,leaflets sessile in 2 – 26 pairs. Inflorescence a few tomany flowered raceme. Bud scales, usually caducous,imbricate, distichous, pale green to brown, glabrous orhairy. Floral bracts persistent or caducous. Bracteolespersistent, petaloid, showy, usually narrow. Hypanthiumnarrowly campanulate, up to 3 mm deep. Sepals 4.Petals 0. Stamens 8 – 12 but usually 10, filaments free.Ovary on a stipe up to 2 mm long, hairy, fused to theinner adaxial wall of the hypanthium; ovules 2 – 3.Fruit a pod, laterally compressed, valves triangular,broadest towards apex, up to c. 10 cm long, woody, 2-valved, glabrous or pubescent, upper suture slightlybroadened, short beak present. Seeds 1 – 2, (whereknown) elliptic or discoid.

Distribution and habitat diversityOf the 18 African phytochoria defined by White(1983), Talbotiella is confined to Phytochorion I, TheGuineo-Congolian region, comprising the Guinean

rainforests which range from Senegal to Gabon andthe Congolian rainforests comprising those of Congo(Kinshasa) together with the bordering forests ofCentral African Republic and Uganda. The Guineanforest is further subdivided into Upper Guinea (Sen-egal to Gabon) and Lower Guinea (Nigeria toGabon). Upper and Lower Guinea are separated bythe more arid Dahomey interval (Togo and Benin).Talbotiella gentii from Ghana is the sole Upper Guineaspecies (Map 1), seven are lower Guinea species; T.eketensis is restricted to SE Nigeria; T. batesii is found insouthern Cameroon, Gabon and Congo Brazzavilleand five species (four of which are described here asnew) are endemic to Cameroon. Two of the newCameroonian species, T. korupensis and T. velutina areendemic to the Caesalpinioideae-rich forest in southKorup, SW Cameroon (Map 1). Several other Caesal-pinioid tree species new to science have beendescribed recently from the Caesalpinioideae-richforest in south Korup (Mackinder & van der Burgt2009; van der Burgt et al. 2007; Wieringa 1999).Although a relatively small genus, the species ofTalbotiella occupy a range of habitats including coastal

Map 1. Distribution of Talbotiella species.


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swamp forest (T. eketensis); lowland rainforest (T.korupensis and T. velutina); savanna-edge, rocky out-crops or rocky plateaus with shallow soil and riverineforest (T. batesii); dry semi-deciduous forest on rockyground (T. gentii); inselberg, rocky outcrops or rockyplateaus (T. batesii, T. breteleri and T. ebo), the latteralso in deciduous forest, and submontane forest (T.bakossiensis). Ostensibly, the genus appears to occupya diverse and disparate assemblage of habitats.However, on closer inspection, there is a generaltrend in the habitats of the majority (five) of thespecies, that of drought resistance. The (brackish)coastal swamps of T. eketensis are physiologically dry.T. gentii grows in the final reaches of the semi-deciduous forest just before the forest yields tosavanna and three more species grow on very shallowsoils of rocky outcrops and rocky plateaus. Only thetwo species from Korup grow in rainforest, while wehave no clear idea of the hydrophysiology of T.bakossiensis. Nevertheless, that at least five out of eightspecies occupy dry forests is noteworthy because themajority of genera in Detarieae occur principally orsolely in wet forest. Only a handful of Africandetarioid genera e.g. Brachystegia, Isoberlinia andJulbernardia show a clear preference for drier habitats,but then the drier habitats are those of savanna and

woodland, not the dry closed forests in whichTalbotiella seems to specialise.

MeasurementsSince it is desirable measurements be comparable, therange of leaflet length and leaflet width given inTable 1 refers to that of the largest pair of leaflets ofthe seven multi-jugate species because leaflet sizevaries considerably depending on the position of theleaflet. An exception was made for the bi-jugatespecies Talbotiella breteleri because the four leaflets areof similar size, so the leaflet size range for this singlespecies is the mean size of all four leaflets. Measure-ment of characters used in the expanded descriptionof T. batesii and in the descriptions of the four newspecies is straightforward but if in doubt, the usershould refer to Wieringa (1999).

Distinguishing speciesWith the exception of the bi-jugate Talbotiellabreteleri, the species of Talbotiella cannot be distin-guished from each other by a single foliage characterbut can be separated by unique combinations ofvegetative characters as presented in Table 1. As

Table 1. Vegetative, distribution and habitat characters distinguishing the species of Talbotiella Baker f. sorted in ascending orderof number of leaflet pairs.

Talbotiellaspecies

No. ofleafletpairs

Leaflet length(mm) Leaflet width (mm)

Leaflet length;presented asapproximatemultiple ofleaflet width Distribution, habitat and altitude

T. breteleri 2 35 – 90 16 – 38 2× CAMEROON: Central.Inselberg, c. 900 – 950 m

T. gentii 5 – 7 15 – 28 8 – 12 2× GHANA: East and Ashanti.Rocky areas in dry semi-deciduous forest,often locally dominant; 100 – 350 m

T. ebo 8 – 11 12 – 34 9 – 11 1 – 3× CAMEROON: Southwest and Littoral.Granitic outcrop in with inselbergvegetation in primary to very oldsecondary, submontane forest andlowland deciduous forest; 100 – 900 m

T. batesii 9 – 13 10 – 15 1 – 3.5 5 – 10× CAMEROON: South. GABON: Woleu Ntem.CONGO-(BRAZZAVILLE): Sangha.Inselberg, rocky outcrops, plateaus orhills, savannah-edge and river banks;500 – 900 m

T. eketensis 12 – 17 9 – 15 2.5 – 4 3.5 – 4× NIGERIA: Southeast.Coastal freshwater swamp forest; nearsea-level

T. velutina 14 – 22 13 – 24 4 – 6 3 – 4× CAMEROON: Southwest.Lowland primary rainforest on well-drained sandy soil, 50 – 100 m

T. korupensis 16 – 22 22 – 35 5 – 9 4× CAMEROON: Southwest.Lowland primary rainforest on well-drainedsoil and on periodically inundated soil;50 – 100 m

T. bakossiensis 21 – 26 12 – 24 2 – 4.5 6× CAMEROON: Southwest.Submontane forest; 850 – 1800 m

403A REVISION OF THE GENUS TALBOTIELLA BAKER F.

restricted distribution and habitat requirements alsocharacterise the species, they have been included inthe table to assist the user in the identification ofspecies.

1. Talbotiella bakossiensis Cheek sp. nov. aff. H. batesiised foliis 7.5 – 13 cm (nec 5.8 – 7 cm) longis, foliisplantarum maturarum 21 – 26 (nec 9 – 13)-jugis,bracteis floralibus dense adpresse tomentosis (necmodice pubescentibus) differt. Typus: Cameroon,South West Province, Bakossi Mts, Mwanzum toKodmin between hut 1 and 2, 18 Nov. 1998, Cheek9688 (holotypus K; isotypi P, WAG, YA).

http://www.ipni.org/urn:lsid:ipni.org:names:77105954-1

Talbotiella bakossiensis ined. Mackinder (2004: 316).

Tree to c. 20 m, dbh up to 40 cm; bole weakly 5-fluted(Cheek 10382), base with small buttresses; bark slightlyrough, covered in mosses; outer slash thick with darkouter line, dull orange, granular-sandy; inner slashpale orange-yellow, fibrous; crown dense. Twigs mod-erately to densely pubescent, hairs 0.5 – 1 mm long.Bud scales 5 – 10, caducous, distichous, brown, coria-ceous, not keeled, outer and inner surface glabrous,proximal scale orbicular, c. 1.5 mm diam., distal scalesbecoming progressively longer and relatively nar-rower, apical scale obovate, 12 × 8 mm, leafy, notkeeled, pubescent along midrib. Stipules (seen inyoung foliage), in pairs, free, auriculate at base, theauricle suborbicular with 2 – 6 acute triangular lobes,up to 0.2 mm long, glabrous on both surfaces butmargins ciliate, upper part of stipule narrowly lanceo-late, 8 – 17 × 1.5 – 2 mm; outer surface denselypubescent along the midrib, veins sparsely pubescent,margins ciliate, inner surface glabrous, apex acute.Leaves alternate, paripinnate, 7.5 – 13 × 1.9 – 3.5 cm;petiole 1 – 2 mm; leaf rachis 6.1 – 11.5 cm long,moderately pubescent, a tuft of longer hairs whereleaflets are inserted; leaflets sessile, in 21 – 26 pairs[18 – 21 pairs in saplings], upper and middle pairsopposite, lower 3 – 5 pairs subopposite; 12 – 24 × 2 –4.5 mm [18 – 22 × 4 – 6 mm in saplings], narrowlyoblong, base asymmetric, proximal margin taperingtowards the apex, glabrous above, lower surfaceappearing glabrous with a hand lens but moderatelyappressed puberulous under a microscope, fringinghairs 0.5 – 0.75 mm long, present particularly on thelower margins on young leaflets but scattered orabsent on mature foliage, midvein central, prominentabove and below for most of its length, becomingobscure just before the apex, small glands 1 – 2 (– 3),not visible with a hand lens but visible under amicroscope, present on the distal half of the lowersurface, positioned about midway between midvein

and margin, always present on basal pair, present orabsent on middle pairs, not seen on uppermost pair.Inflorescence at least a 7-flowered raceme (only a partialinflorescence seen), axis of partial inflorescence c.3 cm long, with a moderate to dense indumentum ofgolden patent hairs, c. 0.6 – 1.5 mm long; floral bractsslightly spathulate, 6 – 10 × 1 – 3 mm, outer surfacedensely appressed tomentose, inner surface glabrous,persistent; pedicels with moderately spreading hairs0.3 – 0.5 mm long; portion of pedicel below bracteoles2 – 4 mm long; bracteoles subopposite to opposite,narrowly oblong, c. 5 × 1.2 – 1.5 mm, tapering towardsthe apex in upper third, mostly glabrous but withmargins ciliate at least in the lower half and along thebasal part of the midvein, a tuft of hairs at the apex;portion of pedicels above bracteoles c. 2 mm.Hypanthium narrowly campanulate, 1 – 1.5 mm deep,puberulous at the mouth, otherwise glabrous on bothsurfaces. Sepals 4, white, subequal, cucullate, 5 – 7 ×2.5 – 3 mm, glabrous on both surfaces except forscattered marginal pubescence on upper third withtuft of hairs 0.2 – 0. 3 mm long at the rounded apex,sessile. Petals absent. Stamens 8, filaments free, 7 – 9 mmlong, glabrous, anthers elliptic, c. 1.2 mm long. Ovary c.2.3 – 2.7 × 1.1 – 1.5 mm, densely golden to red-browntomentose, on a glabrous stipe 1.5 – 2 mm long, fusednear the apex of the hypanthium, free part of stipe veryshort, ovules 2, style c. 3 – 4 mm long, moderately todensely pubescent for most of its length, becomingglabrous just before the stigma, stigma capitate. Pod,seed and seedling unknown. Fig. 1.

DISTRIBUTION. Cameroon, Southwest Province.SPECIMENS EXAMINED. CAMEROON. Southwest Prov-ince: Kupe-Mwanenguba, Nyale, 5°00'N 9°38'E, 18 Nov.1998, Cheek 9678 (K, WAG, YA); Kupe-Mwanenguba,Nyale, Mwanzum to Kodmin between hut 1 and 2,4°59'N 9°40'E, 18 Nov. 1998, Cheek 9688 (K, P, WAG,YA); Kupe-Mwanenguba, Ngomboku. Abang road andthen right to forest, 4°55'N 9°44'E, 11 Dec. 1999, Cheek10326 (K, YA); Kupe-Mwanenguba, Ngomboku, forestridge S of Abang track, 4°55'N 9°44'E, 15 Dec. 1999,Cheek 10382 (K, YA); 35 km NNW Kumba, 2 km SWDikome Balue, Rumpi Mts, Mt Rata, 4°54'N 9°14'E,24 March 1976, Letouzey 14546 (BR, HBG, K, P, WAG,YA); Letouzey 15185 (BR, K, P, WAG); Rumpi Hills, nearDikome (Bakume) Balue, 4°54'N 9°16'E, Nov. 1991,McKey Rh2/43 (K, WAG).HABITAT. Submontane forest; 850 – 1800 m.CONSERVATION STATUS. This species is assessed here asCritically Endangered (CR D) in accordance with thecriteria of IUCN (2001) because less than 50 maturetrees have been found and from only three localities,namely Ngomboku, Mwanzum, and Dikome-Balue.ETYMOLOGY. Named for the Bakossi Mountains, oneof the two areas in Cameroon where the species wascollected and the area where the type was found.




Fig. 1. Talbotiella bakossiensis. A habit; B detail of leaflet; abaxial surface; C habit, stem with leaves flushing; D stipule; E bud-scales; F inflorescence; G floral bract; H flower, side view; J longitudinal section of flower, showing hypanthium and ovary; K detailof anther. A, B, F – K from Cheek 9688, C – E from Cheek 10326. DRAWN BY MARGARET TEBBS.


2. Talbotiella batesii Baker f. (1929: 197). Type:Cameroon, Yaoundé, Bitye, Bates 1596 (holotype BM;isotypes BR, P).

Tree 5 – 12 m, dbh 20 – 60 cm; bark smooth, palegrey, 6 mm thick. Twigs glabrous or sparsely tomoderately pubescent, hairs c. 0.5 mm long. Bud scales3 – 5, caducous; distichous, brown, coriaceous, notkeeled, outer and inner surface glabrous, proximalscale orbicular, c. 1.5 mm diam., distal scales becom-ing progressively longer and relatively narrower, apicalscale obovate, c. 7 × 4 mm, not keeled. Stipules (seenin young foliage) in pairs, free, auriculate at base, theauricle suborbicular, c. 3 × 2 mm, with a singletriangular lobe, c. 1.5 mm long, glabrous on bothsurfaces but margins ciliate, upper part of stipulenarrowly lanceolate, 13 – 15 × 2 – 2.5 mm, outersurface pubescent along the midrib and margins,inner surface glabrous, apex acute. Leaves alternate,paripinnate, 5.8 – 7 × 2 – 2.9 cm; petiole 1 – 2 mmlong, leaf rachis 4.2 – 6.6 cm long, sparsely tomoderately pubescent (most densely within the canalon the distal two-thirds of the rachis, often becomingglabrous in the proximal third), leaflets sessile, in 9 –13 pairs, upper and middle leaflets opposite, lower 2 –3 pairs sub-opposite, 10 – 15 × 1 – 3.5 mm, narrowlyoblong, base asymmetric, proximal margin taperingtowards apex, glabrous above, lower surface appearingglabrous with a hand lens but sparsely appressedpuberulous under a microscope, fringing hairs c.0.5 mm long, present particularly on the lowermargins of young leaflets but absent in mature foliage;midvein central, prominent above and below for mostof its length, becoming obscure just before the apex, asingle small gland, not visible with a hand lens butvisible under a microscope present on the distal halfof the lower surface, positioned about midwaybetween midvein and margin, always present on basalpair, often absent from upper pairs and not seen onthe uppermost pair of leaflets. Inflorescence a 10 – 18-flowered raceme, axis 2.5 – 4.5 cm long including apeduncle c. 2 mm long, light green, with a moderateto dense indumentum of golden patent hairs 1 –1.5 mm long; bud scales pale green; floral bractsobovate to spathulate, 7 – 10.5 × 1.5 – 3 mm, palegreen to white, outer surface moderately pubescent,margins ciliate in upper half but often glabrous inlower half, inner surface glabrous; pedicels white,portion of pedicel below bracteoles 3 – 8.5 mm long,hairs up to 1 mm long; bracteoles opposite, linear tonarrowly oblong, 6 – 8.5 × 1.1 – 2.5 mm, taperingtowards the apex in upper third, white, glabrous orwith a few scattered hairs outside, margins oftenciliate; portion of pedicel above bracteoles 1 – 2 mmlong, scattered spreading pubescence. Hypanthiumnarrowly campanulate, white, 0.7 – 1.3 mm deep,outer surface with scattered hairs in the lower half,

otherwise glabrous, inner surface glabrous. Sepals 4,white, cucullate, outer surface glabrous, inner surfaceglabrous or with scattered erect hairs, 0.3 – 0.7 mmlong around the base, apex with a tuft of ciliate hairsat the margins; adaxial sepal 4 – 5 × 3.5 – 4.5 mm,apex sometimes emarginate, with a tuft of hairs eitherside of the sinus, lateral and abaxial sepals 4 –5.5 × 2.3 – 3.5 mm. Petals absent (rarely a poorlydeveloped linear petal present, 1 × 0.05 mm). Stamens10 (– 12), filaments free, 5 – 9 mm long, white,glabrous or with some scattered hairs, anthers oblong-elliptic, c. 1 mm long, orange. Ovary 1.5 – 3.5 × 1 –1.3 mm, pale pink, white tomentose (golden whendry), on stipe 1 – 2 mm long, fused to the adaxial sideof the hypanthium, free part of stipe very short, c.0.3 mm long, velutinous, ovules 2 – 3, style 4 – 6 mmlong, greenish white, glabrous except for some longhairs at the base, stigma capitate, pale yellow. Podcompressed, 4.2 – 5.5 × 1.8 – 2.7 cm, triangular,broadest towards apex, upper suture slightly broad-ened, 3 – 4 mm wide, surfaces and suture moderatelygolden puberulous, beak 2 – 3 mm long; singledeveloped seed c. 12 × 9 mm, discoid, testa deepreddish brown (Reitsma 3010). Seedling unknown.

DISTRIBUTION. Cameroon, Gabon and Congo-Brazzaville.SPECIMENS EXAMINED. CAMEROON. Bitye, on bank ofthe Ndu, 3°01'N 12°22'E, 14 Feb. 1920, Bates 1596 (BM,BR, P); 33 km S of Djoum, near Akonotangan, close toMiefe R., 2°37'N 12°40'E, 16 Nov. 1966, Letouzey 8409(P). GABON.Minkébé National Park, southern inselbergarea, 1°22'N 12°33'E, 2 May 2003, Ngok Banak et al.1549a (WAG); Rocky outcrop in forest, without trees,c. 25 kmNWofOveng, 0°45'N 11°12'E, 11 Feb. 1987, J. M.Reitsma & B. Reitsma 3010 (LBV, WAG); Ivindo NationalPark, c. 3 kmW of Langoué Bai at WCS camp. Savanna-edge habitat, 0°10'S 12°32'E, 26 Nov. 2002, Stone et al.3486 (LBV, MO, WAG); Mt Sassamongo, rocky plateauW of Sassamongo village, 0°49'N 13°27'E, 9 Jan. 2001,Wieringa et al. 4051 (LBV, WAG). CONGO-BRAZZAVILLE.Sangha, on W slope of Mt Naemba, 1°51'N 13°58'E,15 Nov. 1991, Thomas 8860 (MO, WAG).HABITAT. Inselberg, rocky outcrops or rocky plateaus,savanna-edge and river banks; 500 – 900 m.CONSERVATION STATUS. Using the criteria of IUCN(2001) we here assess this species as CriticallyEndangered (CR D) because it is known from onlysix mature individuals. However, these seven collec-tions are from different localities spread over acircular range of approximately 350 km in diameter.We suspect that a concerted search in areas ofsuitable habitat (inselberg areas) between the knownlocalities might result in the discovery of moreindividuals of Talbotiella batesii. However, since thehabitat type of this species is quite rare, until such asearch has been undertaken, we recommend anassessment of CR D.


ETYMOLOGY. Named for the collector of the type, theAmerican missionary, George Latimer Bates, who col-lected about two-thousand numbers in Cameroon, Equa-torial Guinea and northern Gabon between 1895 – 1928(Letouzey 1968, WAG database).NOTES. At the outset of this study, Talbotiella batesii wasknown only from the type gathering from Cameroon.However, during the course of the work, the authorsfound six additional conspecific collections, one fromCameroon, four from Gabon and one from Congo-Brazzaville. An updated description of T. batesii hasbeen prepared using all the collections.

3. Talbotiella breteleri (Aubrév.) Mackinder & Wieringacomb. nov. Type: Cameroon, Central Province, MtFébé, 3 km NW of Yaoundé, 21 Dec. 1961, Breteler et al.2232 (holotype P; isotypes A, BR, E, FI, K, LISC, SL,UC, WAG, YA).


Hymenostegia breteleri Aubrév. (Aubréville 1970: 104).

Tree, 12 – 22 m, dbh to 80 cm, bole slightlychannelled, bark grey, rough, coarse flakes. Stemspubescent when young, hairs to 0.5 mm long. Budscales up to 9, caducous, distichous, coriaceous, notkeeled, outer surface sparsely hairy, margins ciliate,inner surface glabrous, proximal scale c. 1 mm longand 2 mm wide, distal scales becoming progressivelylonger and relatively narrower, to 7 mm long and4 mm wide, then becoming thinner, first at the edges,and gradually changing to a pair of stipules. Stipules(seen in young foliage) in pairs, free, auriculate atbase, the auricle suborbicular, c. 1.5 × 2 mm, almostglabrous, margin ciliate; upper part of stipule narrowlylanceolate, 13 – 16 × 1.5 – 2 mm, glabrous, marginciliate, apex acute. Leaves alternate, paripinnate, 7 –14 × 5 – 10 cm; petiole 4 – 6 mm, leaf rachis 1.1 –6.2 cm long, canaliculate, pubescent to glabrescent;leaflets sessile, in 2 pairs (sometimes 3 pairs insaplings), rhomboid, asymmetric, apex rounded,upper pair opposite, lower pairs subopposite oropposite, 35 – 90 × 16 – 38 mm, lower pair slightlysmaller than upper pair(s), young leaves appressedpuberulous above and below, with longer hairs onmidrib and margin, mature leaves glabrous above,appressed puberulous below, no fringing hairs, mid-vein central, prominent above and below; small glands0 – 5, not visible with a hand lens but visible under amicroscope, positioned about midway between themidvein and the margin, usually on the distal part ofthe lower surface, sometimes a single gland onproximal part of the lower surface. Inflorescence ac. 10 – 25-flowered raceme; bud scales resembling thoseof the stems, gradually becoming thinner and changing

to floral bracts; inflorescence axis 15 – 45 mm longincluding a peduncle c. 5 – 10 mm long, covered withpatent hairs to c. 1 mm long; floral bracts white,persistent, narrowly oblong, c. 9 × 2 mm, glabrous onboth surfaces, margin ciliate; pedicels covered withpatent hairs to 1 mm long, portion of pedicels belowbracteoles 6 – 9 mm long; bracteoles white, subopposite,narrowly oblong, 8 – 9 × 2 – 2.5 mm, glabrous, marginciliate; portion of pedicel above bracteoles 1 – 2 mmlong. Hypanthium narrowly campanulate, c. 1.5 mmdeep, sparse hairs to 1 mm long on outer surface, innersurface glabrous. Sepals 4, white, dark green at base,opening partly, concave, broadly ovate, outer and innersurface glabrous, apex rounded with a few hairs, adaxialsepal c. 5 × 4 mm, lateral and abaxial sepals c. 5 ×3.5 mm. Petals absent. Stamens 10, filaments white, free,6 – 9 mm long, glabrous; anthers brown, oblong-elliptic,c. 1 mm long. Ovary 2 – 3 mm long, densely covered inpale golden hairs, on a short stipe fused to the adaxialside of the hypanthium; ovules 2; style 5 mm long, sparsehairs on lower third part, glabrous on upper part, stigmacapitate. Pod, seed and seedling unknown.

DISTRIBUTION. Cameroon, Central Province.SPECIMENS EXAMINED. CAMEROON. Central Province:Mt Fébé, 3 km NW of Yaoundé, 3°55'N, 11°29'E,21 Dec. 1961, Breteler 2232 (A, BR, E, FI, K, LISC, P,SL, UC, WAG, YA); hill N of Nkolbisson, 7 km W ofYaoundé, 3°53'N, 11°27'E, 22 Dec. 1961, Breteler 2264(BR, FI, K, LISC, P, WAG, YA); 10 March 2007,Wieringa 5807 (K, WAG, YA).HABITAT. Inselberg; 900 – 950 m.CONSERVATION STATUS. This species is assessed hereas Critically Endangered (CR D) in accordance withthe criteria of IUCN (2001) because less than 50mature trees have been found. Of the three collec-tions known, one is from Mt Fébé where very littlenative vegetation remains (2007 pers. obs.). We believethe other two collections, from a hill north ofNkolbisson (Breteler 2264 and Wieringa 5807), may befrom the same individual but made at an interval ofover 40 years during which time the vegetation on thehill has become very degraded (Breteler pers comm.).ETYMOLOGY. Named for Franciscus (Frans) JozefBreteler, collector of two of the three gatherings fromwhich the species is known.

4. Talbotiella ebo Mackinder & Wieringa sp. nov. affinisH. batesii sed foliis 8.5 – 13.5 cm (nec 5.8 – 7 cm) longis,4.8 – 5.8 cm (nec 2 – 2.9 cm) latis, foliolis 1.25 – 3-plo(nec 5 – 10-plo) longioribus quam latioribus, caulibuspuberulis (nec pubescentibus), parte distali pedicelli(post bracteis locata) glabra vel pilis sparsissimis tanteornata, (nec pilosa) differt. Typus: Cameroon, nearNdokmen II, 8 km E of Yingui, 35 km E of Yabassi, 9 Jan.1972, Letouzey 10940 (holotypus K; isotypi P, WAG, YA).




Tree 15 – 35 m, to 75 cm diam., bole slightly irregular,buttresses up to 2 m high, 1.7 m wide and 15 cm thick;bark smooth, brownish-grey, large flakes sloughing offleaving shallow craters, slash 17 mm thick, fibrous,pinkish-orange, rapidly becoming orange, sticky but noresin seen, sapwood cream-coloured (Wieringa 5880),crown dense, hemi-spherical, reaching down to about4m from the ground, young branches brown, shiny, withfine lenticels. Stems moderately puberulous whenyoung, more sparsely so when older, hairs c. 0.2 mmlong. Bud scales 7 – 11, distichous, persistent orcaducous; brown, coriaceous, not keeled, outer surfacegolden tomentose becoming glabrous at the margins,inner surface densely golden tomentose, proximal scaleorbicular, c. 3 mm diam., distal scales becomingprogressively wider. Stipules (seen in young foliage) inpairs, free, deep-red, soon turning brown and drop-ping, auriculate at base, the auricle suborbicular, c. 3 ×2.5 mm, margins sparsely pubescent, central area ofauricle more densely so; upper part of stipule narrowlylanceolate, 10 – 14 × 2 – 3 mm, midrib moderatelypubescent, otherwise glabrous, apex acute. Leaves alter-nate, paripinnate, 8.5 – 13.5 × 4.8 – 5.8 cm includingthe 2.5 – 3 mm long petiole, [7.8 – 17.2 × 8.4 – 10.6 cmincluding the 1.5 – 3 mm long petiole in saplings]; leafrachis, moderately to densely puberulous in saplingleaves, becoming glabrous or almost so in mature leavesof flowering branches; leaflets sessile, in 8 – 11 pairs,upper and middle pairs opposite, lower 2 – 3 pairssubopposite, 12– 34 × 9 – 11mm [35 – 66 × 11 – 20mmin saplings], oblong, base asymmetric, proximal margintapering towards the apex, glabrous above except forpubescence on lower part of the midvein, lower surfacemoderately to densely and minutely appressed puber-ulous, barely visible with a hand lens but visible whenviewed under a microscope; fringing hairs not seen,midvein central, prominent above and below for most ofits length, becoming obscure just before the apex, smallglands 2 – 5, not visible with a hand lens but present onthe distal half of the lower surface when viewed under amicroscope, positioned slightly (in the basal part of theleaflet) to clearly (in the middle to apical part of theleaflet) closer to the midvein than the margin, seen onall but the uppermost pair of leaflets. Inflorescence a c.18 – 25 flowered raceme, axis 3.2 – 6.6 cm longincluding a peduncle 2 – 3 mm long; axis with amoderate indumentum of golden patent c. 1 mm longhairs; floral bracts 6 – 11 × 2.5 – 5 mm, outer surfacedensely tomentose, margins ciliate, inner surface gla-brous; portion of pedicel below bracteoles 6 – 12 mmlong, sparse spreading weak pubescence, hairs 0.5 –0.75 mm long; bracteoles sub-opposite to opposite,linear, 7 – 9 × c. 1 mm, glabrous, apex acute; portionof pedicel above bracteoles 1.5 – 2 mm long, glabrous oralmost so. Hypanthium narrowly campanulate, 2 – 3 mm

deep, glabrous on both surfaces. Sepals 4, white, slightlycucullate, glabrous on both surfaces except for a tuft ofhairs up to 0.5 mm long at the rounded apex, adaxialsepal 3 – 6.5 × 4 – 5 mm, apex entire, lateral and abaxialsepals 3 – 4.5 × 4 – 5 mm. Petals absent. Stamens 8 – 10,filaments free, 8 – 12 mm long, glabrous, anthersoblong-elliptic, c. 1 mm long, orange. Ovary 4.5 –6 mm long, densely pale golden tomentose, on a stipe1.5 – 2 mm long, velutinous, fused to the adaxial side ofthe hypanthium, free part of stipe very short, ovules 2,style 5 – 7 mm long, moderately pubescent on lowerthird becoming glabrous above, stigma capitate. Podcompressed, triangular, broadest towards apex, 3.8 –7.6 × 2.4 – 3.9 cm, upper suture slightly broadened, c.3 mm wide; surfaces and sutures glabrous. Seedsunknown. Seedling 20 – 30 cm high, first pair of leavesopposite, leaves 6.2 – 12.5 × 4.8 – 6.5 cm, leaflets in 7 –11 pairs, 12 – 45 × 5 – 12 mm. Fig. 2.

DISTRIBUTION. Cameroon. Southwest and LittoralProvinces.SPECIMENS EXAMINED. CAMEROON. Southwest Prov-ince: Loum F.R., 4°43.55N 9°43.45E, 15 Dec. 1999,Mackinder 266 (K, WAG, YA). Littoral Province: 8 km Eof Yingui (= 35 km E of Yabassi), near Ndokmen II, nextto Makombu R. (former Ndol village), 4°32'N 10°22'E, 9Jan. 1972, Letouzey 10940 (K, P, YA); In the proposed EboForest Reserve, 850 m on Dicam Trail from Bekob Camp,4°21.09'N 10°24.94'E, 10 March 2007, Wieringa 5874 (K,WAG, YA); c. 900 m on Dicam Trail from Bekob Camp,4°21.05'N 10°24.94'E, 10 March 2007, Wieringa 5880 (K,WAG, YA) & Wieringa 5881 (K, WAG, YA).HABITAT. Granitic outcrop with inselberg vegetation inprimary to very old secondary, submontane forest andlowland deciduous forest. 100 – 900 m.CONSERVATION STATUS. This species is assessed here asCritically Endangered (CRD) in accordance with the crite-ria of IUCN (2001) because less than 50 mature trees havebeen found, although in the proposed Ebo Reserveseedlings indicating possible regeneration were found. Noinventory exists for Loum F.R. where some logging, albeitlimited has taken place. The Ebo region is poorly knownbotanically and a checklist of plants for the proposed EboReserve will focus efforts on creating a fuller botanicalinventory in this area. This workmay result in the discoveryof additional mature individuals of Talbotiella ebo.ETYMOLOGY. Named for the Ebo area in the LittoralProvince from where four of the five collections fromwhich the species is described, were gathered.

5. Talbotiella eketensis Baker f. (1914: 52); Hutchinson& Dalziel (1928b: 467); Harms (1915: 439); Baker f.(1930: 768); Keay (1958: 467); Keay et al. (1964: 39);Keay (1989: 210). Type: Nigeria, Ibeno, at estuary ofKwa Ibo R., 1912 – 1913, Talbot 3188 (holotype K,isotypes BM, BR, EAH, LISC, WAG).



Fig. 2. Talbotiella ebo. A flowering branch; B seedling; C bark; D stipule; E stipule lower part, inner surface; F stipule lower part,outer surface; G leaflet from base of leaf; H leaflet from middle of leaf; J leaflet from apex of leaf; K gland on lower leaflet surface;L cross-section of leaf-rachis (fresh); M cross-section of leaf-rachis (herbarium collection); N floral bract inner surface; P floral bractouter surface; Q flower; R longitudinal section of flower; S sepal. A, G – K, N – S from Letouzey 10940, B – F from Wieringa 5880,L, M from Wieringa 5874. DRAWN BY HANS DE VRIES.


Shrub, 1 – 6 m, or tree to 15 m, dbh to 10 cm. Stemspubescent, hairs to 0.6 mm long. Bud scales 8 – 10,caducous, distichous, coriaceous, not keeled, outer andinner surface glabrous, margin ciliate, proximal scale c.1 mm long and 2 mm wide, distal scales becomingprogressively longer and relatively narrower, to 9 mmlong and 4 mm wide, then becoming thinner andgradually changing to a pair of stipules. Stipules (seen inyoung foliage) in pairs, free, auriculate at base, theauricle suborbicular, to 1.5 × 2mm; upper part of stipulenarrowly lanceolate, 10 – 17 × 1 – 2 mm, apex acute,outer surface and margins with long hairs, inner surfaceglabrous. Leaves alternate, paripinnate, 4 – 9 × 1.5 –2.5 cm; petiole 1 – 3 mm, leaf rachis 3.5 – 8 cm long,canaliculate, with hairs to 0.5 mm long, becomingglabrescent; leaflets sessile, in 12 – 17 pairs, upper andmiddle pairs of leaflets opposite, lower pairs suboppositeor opposite, 9 – 15 × 2.5 – 4 mm, narrowly oblong, baseasymmetric, proximal margin tapering towards apex,glabrous above except for pubescent midvein on youngleaflets, lower surface with long appressed hairs, fringinghairs to 0.5 mm long, midvein central, impressed above,prominent below; small glands 0 – 2, not visible with ahand lens but visible under a microscope, positionedabout midway between the midvein and the margin,present on the distal part of the lower surface. Inflor-escence a c. 10 – 20-flowered raceme; bud scales c. 6,resembling those of the stems, largest scales to 4 mmlong and 5 mm wide, progressively becoming thinnerand changing to floral bracts; inflorescence axis 20 –40 mm long including a peduncle c. 6 – 10 mm long,covered with patent hairs to 0.6 mm long; floral bractspersistent, narrowly oblong, 5 – 6 × 0.3 – 0.7 mm,glabrous on both surfaces, margin ciliate; pedicelscovered with patent hairs to 0.5 mm long, portion ofpedicels below bracteoles 7 – 10 mm long; bracteolessubopposite, narrowly oblong, 4 – 6 × 0.3 – 0.8 mm,glabrous, margin ciliate; portion of pedicel abovebracteoles 1.5 – 2 mm long. Hypanthium narrowlycampanulate, c. 1.5 mm deep, sparse hairs to 0.5 mmlong on outer surface, inner surface glabrous. Sepals 4,opening partly, concave, broadly ovate, apex rounded,outer and inner surface glabrous, 3.5 – 4 mm long and2 – 2.5 mm wide, adaxial sepal slightly shorter andwider than the lateral and abaxial sepals. Petals absent.Stamens 10, filaments free, 5 – 7 mm long, glabrous,anthers oblong-elliptic, c. 0.8 mm long. Ovary 2 mmlong, densely covered in pale golden hairs, on a stipe c.1 mm long, fused to the adaxial side of thehypanthium, free part of stipe very short, ovules 2, style2.5 – 3 mm long, sparse hairs on lower half, glabrouson upper half, stigma capitate. Pod, seed and seedlingunknown.

DISTRIBUTION. SE Nigeria.SPECIMENS EXAMINED. NIGERIA. Calabar, Eket Distr.,Stubbs Creek F.R. near Unyene, 4°34'N 8°11'E, 10 Jan.

1959, Keay FHI 37716 (K); Southern Nigeria, EketDistr., Ibeno, at estuary of Kwa Ibo R., 4°35'N 7°57'E,1912, Talbot 3188 (BM, BR, EAH, K, LISC, WAG);Southern Nigeria, Degema Distr., 4°45'N 6°46'E, 1914,Talbot 3625 (BM, BR, K, MO, P, WAG).HABITAT. Coastal freshwater swamp forest; near sea-level.CONSERVATION STATUS. Listed by IUCN as Endan-gered (EN B1+2c), due to ongoing clear-cutting,agriculture and development of industrial infrastructureof its forest habitat (IUCN 2007). We reassesses thisspecies here as Critically Endangered (CR D) using thecriteria of IUCN (2001) because it is known from onlythree collections from two localities. The most recentcollection was made c. 50 years ago in 1959.ETYMOLOGY. Named for Eket, one of the two districtsin Nigeria where the species was collected and thedistrict where the type was found.NOTES. Talbotiella eketensis is known only from twofertile Talbot collections and a single sterile collectionfrom Keay. The latter has slightly larger leaves andleaflets that might be confused with those of T. velutinabut the leaflet shape and the locality indicate that T.eketensis is the correct identification. An earlier reportof the presence of T. eketensis in Korup National Parkby Thomas et al. (2003) was based on a misidentifica-tion of his collection 4739. This is one of thecollections on which T. korupensis Mackinder &Wieringa is based.

6. Talbotiella gentii Hutch. & Greenway (in Hutchinson& Dalziel 1928a: 382); Hutchinson & Dalziel (1928b:467); Baker f. (1930: 769); Keay (1958: 467); Irvine(1930: 403), (1961: 317); Hall & Swaine (1981: 301);Hawthorne (1990: 200). Type: Ghana, Kwahu, NorthScarp, Dec. 1926, Gent 184 (holotype K; isotypes BR,FHO).

Tree, 9 – 18m, dbh to 50 cm, bark smooth or rough, verythin. Stems pubescent, hairs to 0.3 mm long. Bud scalesup to 10, caducous, distichous, coriaceous, not keeled,outer and inner surface glabrous, margins ciliate,proximal scale c. 1 mm long and 2 mm wide, distalscales becoming progressively longer and relativelynarrower, to 7 mm long and 5 mm wide, thenbecoming thinner and gradually changing to a pair ofstipules. Stipules (seen in young foliage) in pairs, free,auriculate at base, the auricle suborbicular, c. 1.5 ×1.5 mm; upper part of stipule linear, 12 – 15 × 1 –1.5 mm, apex acute; outer surface sparse hairy, innersurface glabrous, margin ciliate. Leaves alternate, pari-pinnate, 4 – 10.5 × 2.2 – 6.8 cm; petiole 2 – 5 mm, leafrachis 3 – 7.5 cm long, canaliculate, pubescent, leafletssessile, in 5 – 7 pairs, rhomboid, asymmetric, apexrounded, upper and middle pairs opposite, lower pairssubopposite or opposite, (8 –) 15 – 28 (– 40) × (4 –)8 – 12 (– 16) mm, glabrous above except for pubescent


midvein on young leaflets, appressed puberulous below,fringing hairs to 0.5 mm long, particularly on themargins of young leaflets; midvein running diagonally,prominent above and below; small glands 0 – 3, notvisible with a hand lens but visible under a microscope,positioned about midway between the midvein and themargin, on the distal part of the lower surface.Inflorescence a c. 6 – 12-flowered raceme; bud scalesresembling those of the stems, gradually becomingthinner and changing to floral bracts; inflorescenceaxis red, 21 – 38 mm long including a peduncle c. 2 –13 mm long, covered with patent hairs to c. 0.7 mmlong; floral bracts caducous, narrowly oblong, c. 7 –10 × 1.5 mm, glabrous on both surfaces, margin ciliate;pedicels covered with patent hairs to 0.5 mm long,portion of pedicels below bracteoles 7 – 11 mm long;bracteoles subopposite, linear, 7 – 10 × 0.2 – 0.3 mm,glabrous, margin ciliate; portion of pedicel abovebracteoles 1 – 2.5 mm long. Hypanthium narrowlycampanulate, 2.5 – 3 mm deep, sparse hairs to0.3 mm long on outer surface, inner surface glabrous.Sepals 4, white to pink, opening partly, concave, broadlyovate, outer surface glabrous, inner surface with a fewlong hairs, adaxial sepal c. 5 × 3.5 mm, apex roundedand shallowly emarginate with a few long hairs eitherside of the sinus, lateral and abaxial sepals c. 5 × 3 mm,apex rounded with a few long hairs. Petals usuallyabsent, occasionally a rudimentary petal of 2 × 0.1 mmpresent. Stamens 10, filaments free, 8 – 9 mm long,glabrous; anthers oblong-elliptic, 1.2 – 1.5 mm long.Ovary 2 – 2.5 mm long, white, densely covered in whitehairs, on a stipe c. 1 mm long, fused to the adaxial side ofthe hypanthium, free part of stipe very short; ovules 2;style 6 – 8 mm long, sparse hairs on lower third part,glabrous on upper part, stigma capitate. Pod glossy, lightbrown, triangular, broadest towards apex, 3.5 – 5.5 ×2.2 – 2.5 cm, beak to 3 mm long, upper suture slightlybroadened, 2 mm wide; valves curling when dry; pedicel13 – 20mm long. Seed elliptic, 14 – 17 × 12 – 14 × 3 mm.Seedling : hypocotyl 3 – 4.5 mm long, epicotyl 4.5 – 8.5cm long; first pair of leaves opposite, leaves 3.5 – 6 × 3 –5 cm, leaflets in 3 – 4 pairs, 16 – 30 × 7 – 13 mm,hypocotyl 3 – 4.5 mm long, epicotyl 4.5 – 8.5 cm long.

DISTRIBUTION. Ghana, Eastern & Ashanti Region.SPECIMENS EXAMINED. GHANA. Eastern Region:Krobo Mt, 6°05'N 0°03'E, 15 Nov. 1956, Adams s.n.(GC); 8 Sept. 1952, Boughey s.n. (GC); Akosombo, 6°18'N 0°03'E, 7 Jan. 1974, Enti R1131 (K, MO, WAG); 8Jan. 1974, Enti FE1333 (BR, MO, P, WAG); 25 May1970, Enti & Hall GC40245 (GC); Mpraeso Distr.,Afram Mankrong Forest Reserve, 6°39'N 0°20'W, Nov.1956, Enti FH 6512 (B, K, P); Kwahu Nteso to Tarkwa,6°39'N 0°36'W, 16 Jan. 1968, Enti & Hall GC37476(FHO, K, P); Kwahu Nteso to Ankoma road, Asuboni,6°38'N 0°39'W, 17 Jan. 1968, Enti & Hall GC37490(GC); N end of Krobo Hill, 6°05'N 0°03'E, 4 Jan. 1974,

Faden et al. 74/9 (BR, GC, K, MO); North Scarp,Kwahu, 6°42'N 0°45'W, Dec. 1926, Gent 184 (BR, FHO,K); Summit of Krobo Mt, 6°05'N 0°03'E, 28 Nov. 1958,Hall 1228 (GC); Near Worobong R. on Mpaamu rd,Worobong Forest Reserve, 6°37'N 0°25'W, 12 June1970, Hall & Agyakwah GC39682 (GC); Near Aketswia,Aketswia, 6°32'N 0°12'W, 11 June 1970, Hall & EntiGC39652 (GC); Ashanti Region, Ashanti, Agogo, 6°48'N 1°05'W, Dec. 1927, Irvine 871 (FHO, GC); nolocality, undated, Irvine 957 (FHO, K); Krobo ForestReserve 5 miles above Odumasi, 6°11'N 0°01'E, 24Dec. 1938, Irvine 3072 (GC, P); near Akosombo, 6°18'N 0°03'W, 30 Nov. 1993, Jongkind et al. 1327 (WAG);Krobo Hill, 6°05'N 0°03'E, 8 Dec. 1994, Jongkind et al.1940 (MA, MO, WAG); Volta R. Reserve, 6°09'N 0°04'E, July 1927, Moor 75 (FHO); Volta R. Reserve, 6°09'N 0°04'E, Nov. 1927, Moor 1138 (FHO); Fringingforest above Ajena by Volta R. Ajena, 6°19'N 0°05'E, 28Nov. 1952, Morton GC8030 (GC, K, W, WAG); 3 milesabove Ajena, 6°22'N 0°05'E, 29 Nov. 1953, Morton GC9454 (BH, FHI, GC, K, P); No locality, undated, C. J.Taylor s.n. (BR); No locality, 1954, C. J. Taylor s.n. (BR);No locality, 1956, C. J. Taylor s.n. (BR); Huhunia acc. toF.W.T.A, Vtuhunia on label K, 6°10'N 0°11'W, 15 Jan.1930, A. S. Thomas D57 (GC, K, P); Abetifi, 6°40'N 0°45'W, 10 June 1908, Thompson 70 (GC); No locality, 6°40'N 0°45'W, 12 June 1908, Thompson 87 (K); Akatui,6°31'N 0°09'W, Oct. 1935, Vigne FH4026 (GC); YongwaForest Reserve, 6°14'N 0°03'W, June 1937, Vigne FH4398 (BM, BR, FHO).HABITAT. Dry semi-deciduous forest, on rocky ground,often locally dominant, sometimes forming purestands, often forming a belt between savanna anddeciduous rainforest; 100 – 350 m.CONSERVATION STATUS. Talbotiella gentii is the specieswith the highest conservation priority in Ghana and islisted as Critically Endangered (CR A1c, B1+2c) asassessed by Hawthorne (IUCN 2007) who reports thathabitat loss and degradation due to clear-cutting,agriculture and human settlement are ongoing threatsto this species. T. gentii now exists in small stands inrocky areas of dry forest, most of the original area ofwhich has been extensively destroyed.ETYMOLOGY. Named for the collector of the typespecimen.NOTES. Talbotiella gentii is the best known species of thegenus but was erroneously reported as being presentin Cameroon by Hall (1976) based on a misidentifica-tion of Letouzey 11626 which is instead Hymenostegia cf.floribunda (Benth.) Harms.

7. Talbotiella korupensisMackinder & Wieringa sp. nov.affinis H. eketensis sed foliis 8.5 – 19.5 cm (nec 3.5 –8.7 cm) longis, foliis plantarum maturarum 16 – 22(nec 12 – 19)-jugis, foliolis 22 – 35 × 5 – 9 mm (nec8 – 14 × 2 – 4 mm), bracteis floralibus 2.5 – 3 mm


(nec c. 1 mm) latis, parte proximali pedicelli (infrabracteas) 2 – 4 mm (nec 7 – 12 mm) longis, bracteolisellipticus vel ovatus (nec linearis), 2-plo tantum (nec6 – 8-plo) differt. Typus: Cameroon, Southwest Prov-ince, Korup National Park, near P transect, northeast ofNW plot, subplot 35 IN, 5°01'N 8°47'E, 28 March 2003,van der Burgt & Bischoff 629 (holotypus WAG; isotypi B,BR, FHO, G, K, MA, MO, P, SCA, US, YA).


Tree 8 – 20 m, dbh 15 – 35 (– 65 cm), bole oftensloping or bent, cylindrical to irregular, usually withstem shoots, bark yellowish light brown, smooth but alittle irregular, sometimes flaky. Stems moderatelypuberulous or pubescent, hairs c. 0.2 – 0.6 mm long.Bud scales up to 7, caducous, distichous, medium brown,coriaceous, not keeled, outer surface sparsely hairy,with scattered puberulous margins, inner surface gla-brous, proximal scale orbicular, c. 2.5 mm diam., distalscales similar but becoming progressively longer andrelatively narrower, outer surface becoming progres-sively more densely puberulous. Stipules (seen in youngfoliage) in pairs, free, auriculate at base, the auriclesuborbicular, c. 5 × 3 mm, outer surface veins andmargins sparsely pubescent; upper part of stipulenarrowly lanceolate, 12 – 14 × 3 – 4 mm, outer surfacesparsely hairy, inner surface glabrous, apex acute. Leavesalternate, paripinnate, 8.5 – 19.5 × 3.4 – 5.4 cm; petiole3 – 7 mm, leaf rachis 7.4 – 19.2 cm long, sparsely tomoderately pubescent (most densely so within thecanal on the distal two-thirds of the rachis, becomingalmost glabrous in the proximal third), leaflets sessile, in16 – 22 pairs, upper and middle leaflets opposite, lowerpairs subopposite or opposite, 22 – 35 × 5 – 9 mm,narrowly oblong, base asymmetric, proximal margintapering towards apex, glabrous above except forpubescent midvein on young leaflets, lower surfaceminutely appressed puberulous, fringing hairs 0.5 –1 mm long, particularly on the lower margins of youngleaflets but scattered or absent in mature foliage,midvein central, prominent above and below for mostof its length, becoming obscure just before the apex,small glands 1 – 4, not visible with a hand lens butvisible under a microscope, present on distal part of thelower surface, positioned about midway between themidvein and the margin, seen on all but the upper twopairs of leaflets. Inflorescence a c. 10 – 30-floweredraceme, axis 5.4 – 7. 7 cm long including a peduncle2 – 7 mm long, moderately covered with pale patenthairs c. 1 mm long; floral bracts narrowly oblong, c. 8 ×2 mm, glabrous on both surfaces, caducous; pedicelssparse to moderate spreading pubescence, hairs c.0.5 mm long, portion of pedicel below bracteoles 8 –13 mm long; bracteoles subopposite, elliptic to ovate,5 – 7 × 3 – 4 mm, glabrous, apex rounded; portion ofpedicel above bracteoles 3 – 4 mm long. Hypanthium

narrowly campanulate, c. 2 mm deep, sparsely longpubescent hairs 0.75 – 1 mm long on outer surface,inner surface glabrous. Sepals 4, white, opening onlyslightly, concave, broadly ovate, outer surface long-pubescent on basal half to two thirds, glabrescenttowards margins and on apical third, inner surfaceglabrous, adaxial sepal c. 4 – 5 × 4 mm, apex roundedand shallowly emarginate; with a tuft of hairs either sideof the sinus, lateral and abaxial sepals 4 – 5 × 3 mm,apex rounded with tuft of hairs. Petals absent. Stamensusually 10, filaments free, 7 – 8 mm long, glabrous,anthers oblong-elliptic, c. 1.5 mm long, reddish. Ovary2 – 3 mm long, densely pale golden sericeous, on astipe 1 – 1.5 mm long, fused to the adaxial side of thehypanthium, free part of stipe very short, ovules 2, style4 – 6 mm long, moderately pubescent on lower thirdbecoming glabrous above, stigma capitate. Pod com-pressed, triangular, broadest towards apex, 6.8 – 9.6 ×3.5 – 4.6 cm, upper suture slightly broadened, 2 –3 mm wide; surfaces and suture glabrous. Seedsunknown. Seedling: first pair of leaves opposite, leaves8 – 10.5 × 4– 4.3 cm, leaflets in 10 – 11 pairs, 22 – 25 ×8 – 9 mm. Figs 3 & 4.

DISTRIBUTION. Cameroon: Southwest Province. KorupNational Park and Besingi village forest.SPECIMENS EXAMINED. CAMEROON. Southwest Prov-ince: Korup National Park, 30 mN of Science Camp,5°00'N 8°48'E, 29 May 2000, van der Burgt 601 (G, SCA,WAG); Korup National Park, NE of NW Plot, subplot35 IN, 5°01'N 8°47'E, 28 March 2003, van der Burgt &Bischoff 629 (B, BR, FHO, G, K, MA, MO, P, SCA, WAG,US, YA); Korup National Park, P plot, subplot 23H, 5°01'N, 8°47'E, 8 Sept. 2004, van der Burgt 708 (BR, G, K,MO, P, SCA, WAG, YA); Korup National Park, near Ptransect, northeast of NW plot, subplot 35 IN, 5°01'N8°47'E, 11 Oct. 2005, van der Burgt 770 (BR, G, K, MO,P, SCA, WAG, YA); near Besingi village, upstream ofhigh bridge over Idu R., 4°55'N, 8°54'E, 13 Sept. 2006,van der Burgt et al. 845 (K, MO, P, WAG, YA); KorupNational Park, 5°03'N 8°48 E, 28 Feb. 1984, D. W.Thomas 3252 (K, MO, P, WAG, YA); between Bulu andDibunda, 4°54'N 8°53'E, 16 Dec. 1984, D. W. Thomas4155 (MO); Korup National Park, between Ndian riverat PAMOL field and 2.5 km on transect "P", 5°01'N 8°50'E, 12 April 1985, D. W. Thomas 4739 (B, BR, K, LE,MO, P, S, WAG).HABITAT. Lowland primary rainforest, on well-drainedsoil and on nearby periodically inundated soil; 50 –100 m. The type locality of Talbotiella korupensis is inthe “P transect” plots, in the southern part of KorupNational Park. These plots have a total size of155.75 ha. All trees of over 50 cm trunk diameterwere registered and identified. Of the 3181 recordedtrees, one was identified as T. korupensis. The trunkdiameter of trees of this species is usually less than35 cm. Trees with a trunk diameter of between 10 and



Fig. 3. Talbotiella korupensis. A flowering branch;B leaflets from base of leaf, from middle of leaf & from apex of leaf; C cross-sectionof leaf-rachis (herbarium collection); D stipule; E stipule lower part, inner surface; F section of underside of leaflet; G seedling; Hflower; J longitudinal section of flower; K floral bract inner surface; L fruit. From van der Burgt & Bischoff 629. DRAWN BY HANS DE VRIES.


50 cm were recorded in 56 randomly located subplotsof 0.25 ha each; total size 14 ha. Of the 5755 recordedtrees between 10 and 50 cm trunk diameter, 44 wereidentified as T. korupensis. The 44 trees were notspread evenly over the subplots; one of the subplotshad nine T. korupensis trees, out of a total of 127 treeswith a trunk diameter of over 10 cm. T. korupensisalways grows gregariously; the species is abundant inmany parts of the south Korup forest but completelyabsent in other parts that otherwise have a similarspecies composition.

The trunk girth of 30 trees was measured twice, at aheight of 1.3 m, with an interval of 16.3 to 16.9 years.The average trunk diameter increment of the 30 treeswas 0.9 mm/year.CONSERVATION STATUS. Talbotiella korupensis isassessed here as Endangered (EN D) under thecriteria of IUCN (2001). This species is locallyabundant in south Korup and also recorded fromthe forests of Besingi village.ETYMOLOGY. Named for Korup National Park wheremost of the specimens from which the species isdescribed were gathered, and which holds the mostimport stands of this species.NOTES. Trees of Talbotiella korupensis can be difficult todistinguish from T. velutina when not in flower. Bothspecies have only been found in the southern part ofKorup National Park, and in the forest of Besingivillage, at 15 – 18 km to the southeast of the type

locality. Sometimes trees of the two species grow nextto each other. Juvenile trees of the two species areespecially difficult to separate. Comparing leaflet sizemay be the best way of separating juvenile trees(Table 1). Adult trees can be separated by comparingsimultaneously the leaflet size, size of the trees, theirstem shape, and the habitat type. T. korupensis is anunderstory tree, while T. velutina is a canopy tree.Adult trees of T. korupensis are usually 5 – 20 m highand have a 10 – 35 cm trunk diameter; while adulttrees of T. velutina are usually 20 – 35 m high and havea trunk diameter of up to 76 cm. Trees of T. korupensiscan have vertical trunks but they are often sloping,cylindrical to irregular, and stem shoots are almostalways present; while the trunks of T. velutina arealways vertical, fluted or lobed up to the crown, andusually without stem shoots. T. korupensis is usuallyfound on or near periodically inundated soil, while T.velutina is only found on well-drained soil.

8. Talbotiella velutina Burgt & Wieringa sp. nov. affinisT. korupensis sed trunco lobato vel sulcato (neccylindrico) surculis plerumque nullis (nec saepepraesentibus), axe inflorescentiae 1.6 – 5.2 cm (nec4.5 – 8.5 cm) longo indumento densissimo velutinoaureo axem omnio obtegenti nec axe pilos pallidospatentes aliquantum crispos modice tantum abun-dantes ferenti, bracteis floralibus extus modice vel

Fig. 4. Talbotiella korupensis. A flowering branch; B base of a tree 27.8 cm diameter at 1.4 m. A from van der Burgt & Bischoff629, B not collected. PHOTOS: A WOLFGANG BISCHOFF; B XANDER VAN DER BURGT.


dense adpresse pubescentibus (nec glabris), 3 ½ vel 4-plo (nec usque duplo tantum), legumine modicepuberulo (glabro) differt. Typus: Cameroon, KorupNational Park, PE plot near P transect, subplot 16ZN,tree number PE8123, 5˚01'N 8˚48'E, alt. c. 100 m,15 Oct. 2005, van der Burgt & Eyakwe 777 (holotypusWAG; isotypi B, BR, G, K, MA, MO, P, SCA, US, YA).


Tree 26 – 35.3 m, dbh 22 – 76 cm; bole straight,vertically lobed or fluted along its entire length,sometimes with steep buttresses, up to 2.5 m highand c. 1 m wide; bark smooth, pale light brown. Stemsmoderately pubescent, hairs to c. 1.5 mm long. Budscales up to 15, distichous, caducous, coriaceous, notkeeled, outer surface moderately puberulous, at leasttowards the edges, margins ciliate, inner surfaceglabrous, light brown-green with reddish edges (whenfresh), uniformly brown when dried, proximal scalessuborbicular, c. 1.5 – 2 mm diam., distal scalesbecoming progressively longer and relatively nar-rower, on branches in leaf flush apical scale obovate,c. 23 × 18 mm, not keeled; on inflorescence branchapical scale to 9 × 10 mm. Stipules (seen in youngfoliage) in pairs, free, auriculate at base, the auriclesuborbicular, to 6 × 3 mm, margins ciliate, upper partof stipule narrowly lanceolate, 15 – 20 (– 28) × 2 (– 3)mm, apex acute, outer surface and margins appressedpubescent, inner surface glabrous. Leaves alternate,paripinnate, 5.2 – 11 (– 17.5) × 2.4 – 4.3 (– 5.1) cm;petiole 2 – 4 mm long, leaf rachis 4.4 – 9.8 (– 15.6)cm long, sparsely to moderately pubescent, leafletssessile, in 14 – 22 pairs, upper and middle leafletsopposite, lower 3 – 4 pairs sub-opposite, (7 –) 13 – 24(– 32) × (2 –) 4 – 6 (– 8) mm, narrowly oblong, baseasymmetric, proximal margin tapering towards apex,apex rounded, glabrous above except for pubescentmidvein, lower surface sparsely appressed puberulous,barely visible with a hand lens but visible under amicroscope, fringing hairs c. 2 mm long, on the lowermargins of young leaflets but absent from seedlingand mature leaflets; midvein central, prominent aboveand below for most of its length, becoming obscurejust before the apex, in leaflets 0 – 4 (– 7) small glandspresent, not visible with a hand lens but visible with amicroscope, on the distal half of the lower surface,commonly positioned about midway between themidvein and margin. Inflorescence an 8 – 20-floweredraceme, axis 16 – 52 mm long including a peduncle8 – 15 mm long, peduncle and rachis with a denseindumentum of golden sericeous hairs, 1 – 1.5 mmlong. Floral bracts often caducous, obovate to spathu-late, 7 – 10.5 × 1.5 – 3 mm, outer surface moderatelyto densely appressed pubescent, upper margins ciliatebut margins often glabrous in the lower half, innersurface glabrous; pedicels white, moderately to densely

crinkly patent hairs to 1.2 mm long, portion of pedicelbelow bracteoles 6 – 8 mm long; bracteoles white,opposite, linear to narrowly elliptic, 6 – 8 × 1 – 2 mm,densely appressed pubescent outside, glabrous inside,margins ciliate; portion of pedicel above bracteoles 3 –4 mm. Hypanthium narrowly campanulate, white,1.5 mm deep, outer surface moderately patent pubes-cent, inner surface glabrous. Sepals 4, white, cucullate,outer surface and margins sparsely and unevenlypubescent, inner surface glabrous, apex rounded;adaxial sepal c. 4.5 – 6 mm, lateral sepals c. 3 ×6 mm, abaxial sepal c. 4 × 6 mm. Petals absent. Stamens10, filaments free, 8 – 10 mm long, white, glabrous orwith some scattered hairs, anthers oblong-elliptic,1.2 – 1.5 mm long, orange-brown. Ovary white, 2 –3 × 1 mm, very densely covered with spreading goldenpubescent hairs to 1.5 mm long, on a stipe 1 – 2 mmlong, velutinous, fused to the adaxial side of thehypanthium, free part of stipe very short, less than0.5 mm long, ovules 2, style 5 – 7 mm long, sparse longhairs on lower half, stigma capitate. Infructescence axis16 – 30 mm long, retaining dense velutinous indumen-tum, 1 or 2 pods, usually at the top of the axis subtendedby fruiting pedicels 11 – 15 mm, unevenly sparsely tomoderately spreading pubescent. Pod compressed, 4.8 –8.9 × 2.4 – 3.9 cm, triangular, broadest towards apex,upper suture slightly broadened, 4 – 5mmwide, surfacesand suture, moderately puberulous, beak 1 – 2mm long.Seeds 1 – 2 (0 in aborted pods); 1 – 1.5 cm diam., c.3 mm thick. Seedling : hypocotyl 4 – 6 cm, epicotyl 6 –7.5 cm, first pair of leaves opposite, leaves 7.2 × 3.3 cm,leaflets in 12 pairs to 17 × 6 mm. Figs 5 & 6.

DISTRIBUTION. Cameroon, Southwest Province, KorupNational Park and Besingi village forest.SPECIMENS EXAMINED. CAMEROON. Southwest Prov-ince: Korup National Park, P transect, P plot, subplot29A, tree number P6319, 5˚01'N 8˚47'E, pods andseedlings, 8 June 2000, van der Burgt 610 (G, SCA,WAG); near P plot, subplot 31PN, 2 March 2000, vander Burgt 619 (SCA, WAG); P plot, subplot 28B, treenumber P6303, 13 March 2004, van der Burgt, Praz &Eyakwe 663 (BR, G, K, MO, P, SCA, WAG, YA); P plot,subplot 28B, tree number P6303, 13 Sept. 2004, vander Burgt & Eyakwe 712 (BR, G, K, MO, P, SCA, WAG,YA); PE plot near P transect, subplot 16ZN, treenumber PE8123, 15 Oct. 2005, van der Burgt & Eyakwe777 (B, BR, G, K, MA, MO, P, SCA, US, WAG, YA); PEplot near P transect, subplot 16ZN, tree numberPE8123, flush leaves, 26 Oct. 2005, van der Burgt &Eyakwe 787 (BR, G, K, MO, P, SCA, WAG, YA); PE plotnear P transect, subplot 16ZN, tree number PE8123,young fruits, 5 Dec. 2005, van der Burgt & Motoh 814 (K,SCA, WAG); near P plot, subplot 31QN, 6 March 2007,van der Burgt et al. 906 (K, SCA, WAG, YA); near villageBesingi, 4°55'N 8°54'E, seedlings, 12 March 2007, vander Burgt et al. (K, SCA, WAG, YA); near village Besingi,



Fig. 5. Talbotiella velutina. A branch with inflorescences and flushing leaves; B flower; C flower with sepals removed; D infructescencewith two pods; E dried pod valve; F leaf upper surface; G leaflet lower surface; H stipule. A – C, F, G from van der Burgt & Eyakwe 777; Dfrom van der Burgt & Motoh 814; E from van der Burgt & Eyakwe 712; H from van der Burgt & Eyakwe 787. DRAWN BY XANDER VAN DER BURGT.


4°55'20''N 8°53'58''E, seedlings, 15 Feb. 2008, Pearce etal. 5 (BR, G, K, MO, P, SCA, WAG, YA).HABITAT. Lowland primary rainforest on well-drainedsandy soil; 50 – 100 m. The rainfall at the Bulu

weather station, c. 12 km southeast of the type locality,ranged from 4023 to 6146 mm/y, and averaged5029 mm/y (1984 – 2004). The climate is stronglyseasonal with one distinct dry season from December

Fig. 6. Talbotiella velutina. A inflorescence; B flower; C flushing leaves; D base of a tree 54.3 cm diameter at 1.3 m. A – C from vander Burgt & Eyakwe 777. D not collected. PHOTOS XANDER VAN DER BURGT.


to February (average monthly rainfall less than100 mm).

Talbotiella velutina occurs in Caesalpinioideae-richforest and was discovered in and near the “P transectplots” in the southern part of Korup National Park.The total size of these plots is 155.75 ha. Of the 3181recorded trees ≥ 50 cm trunk diameter, eight treeswere identified as T. velutina. Trees with a trunkdiameter of between 10 and 50 cm were registeredin 56 randomly located subplots of 0.25 ha each, atotal size of 14 ha. Of the 5755 recorded treesbetween 10 and 50 cm trunk diameter, none wereidentified to T. velutina. Around the eight trees oftrees ≥ 50 cm trunk diameter, but outside thesubplots referred to above, some trees smaller than50 cm diameter were found. Several groups of treeswere found near the plots (Map 2). In a forest nearthe village Besingi, 15 – 18 km from the other trees, agroup of four trees was found. In total 39, trees ofT. velutina were found.

The trunk girth of six trees was measured twice,with an interval of 14 years (three trees) or 7.2 years(three trees). The average stem diameter increment ofthe six trees was 2.9 mm/year.

Trees of Talbotiella velutina were found flowering inMarch and October. The main flowering period isMarch, when the ground beneath the trees may becovered with fallen flowers; and fruiting occurred inMay and June of the same year with newly fallenpod valves. The trees also flowered once in October,but the flowering was much less abundant, and didnot result in any seeds although a few empty podswere found. The flowers of an inflorescence allopen at the same time and point in the samedirection (Fig. 6A).

The seeds of Talbotiella velutina, as is the case forother species of Talbotiella where fruits are known,are dispersed from the fruits in the tree by ballisticseed dispersal. This is indicated by the fact that thedrying valves of a pod curl in opposite directions(Fig. 5E). The maximum ballistic dispersal distance isunknown, but is estimated to be in the range of20 – 30 m.

Talbotiella velutina occurs in small groups (Map 2).The tendency of the species to grow in groups isprobably related to the limited possibilities of theballistic seed dispersal method. Data of comparabledetail is lacking for the other species. The distancebetween a group of trees and the nearest next groupis c. 400 – 500 m. This indicates the likelihood ofsecondary seed dispersal across distances in thisrange although the method of secondary seeddispersal is not known.

Talbotiella velutina can be difficult to distinguishfrom T. korupensis. See the section on T. korupensis forsome notes on how to distinguish the two species inthe field.CONSERVATION STATUS. Talbotiella velutina is hereassessed as “Critically endangered, CR D” (IUCN2001). In total 39 trees of T. velutina were found; mostof them were mature. The species should be classifiedas Critically Endangered because less than 50 maturetrees have been found.ETYMOLOGY. The epithet ‘velutina’ refers to thedensely hairy inflorescence axis which characterisesthe species.NOTES. Talbotiella velutina and T. korupensis have arestricted and more or less common distribution,sometimes even growing side by side in the forest.Furthermore, they are morphologically closer toeach other than any other known species of Talbo-tiella. These observations lead us to speculate thatthey may represent an example of sympatric speci-ation, i.e. species that have differentiated from acommon ancestor in the absence of a geographicalboundary.

In a phylogenetic analysis of rbcL nucleotidesequence data (Mackinder et al. 2010), all four ofthe sampled species of Talbotiella are placed togetherin a clade with two species of Hymenostegia s.l. but theclade is without internal resolution, thus neithersupporting nor refuting our conjecture concerningthem being sister species, but given the morpholog-ical resemblance, this is at present a most likelyscenario. A combined analysis of nucleotide sequencedata from the rbcL, trnL and matK cpDNA regions isplanned pending the generation of a comparablematK and trnL data set which subsequently may shedfurther light on the derivation of these two species. Ifthey indeed prove to be sister species, the pair couldprovide an interesting case study of potential sympatricspeciation.

Map 2. Talbotiella velutina. Distribution of 35 trees in primaryrainforest in Korup National Park. The species occurs in smallgroups. The area on the map is only partly surveyed; moretrees and groups of trees may be present. The size of a dotrepresents the stem diameter in 7 size classes of 10 cm: 10 –

20 cm up to 70 – 80 cm. The circle represents a dead tree thatbecame uprooted in 2001 or 2002.


AcknowledgementsThe first two authors received support from theSYNTHESYS Project http://www.synthesys.info/ whichis financed by European Community Research Infra-structure Action under the FP6 "Structuring theEuropean Research Area" Programme. Specifically,The Netherlands Taxonomic Facility of the SYN-THESYS project provided funds which allowed thefirst author to travel to the Netherlands to undertake acollaborative study of the generic limits of Talbotiella(NL-TAF-173). We are grateful to the curators of B,BM, BR, FHO, P, MO and YA for the loan of theirspecimens and to Yvette Harvey who, with the kindpermission of the curators of GC, photographed theirholdings of Talbotiella. We would like to thank Jean-Michel Onana and our other colleagues at theNational Herbarium of Cameroon, Yaoundé for theirco-operation and support in Cameroon. We also thankthe villagers of Kodmin who kindly assisted theexploration for T. bakossiensis.

Some of the relevant fieldwork for the project wasdependent on the Earthwatch Institute (Europe) withthe support of DGVIII of the European Commission.The discovery of Talbotiella velutina resulted fromwork that was supervised by Prof. D. M. Newbery ofthe Institute of Plant Sciences, Bern and funded bythe Swiss National Science Foundation. We thank theCameroon government for permission to carry outresearch in Cameroon and Korup National Park(Conservator, A. Kembou) for access to the site.Moses Elangwe climbed the tree from which thetype of T. velutina was collected. Wolfgang Bischoff,Moses Eyakwe and Motoh Jackson provided fieldassistance.

Under the aegis of CRES, with considerable logisticsupport from Bethan Morgan, Head of the Ebo ForestResearch Station and her staff, we were able to visit theproposed Ebo National Park, and locate a secondmature specimen and first stipules and seedlings ofTalbotiella ebo, a species previously only collected oncebefore by Renée Letouzey.

We are grateful to Melanie Thomas for translatingthe diagnoses into Latin. We would also like to thankMargaret Tebbs for preparing the illustration ofTalbotiella bakossiensis and Hans de Vries for preparingthe illustrations of T. ebo and T. korupensis.

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Documents

A revision of the genus Talbotiella Baker f. (Caesalpinioideae: Leguminosae)