Copyright (c) American Society for Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 DOI 10.1600/036364411X553108 © Copyright2011 bythe American SocietyofPlantTaxonomists Systematic Botany restricted distributions. cies are relatively widespread, whiletheotherthree have primary rainforests tosavannas andwoodlands.Two spe- tributed across tropical Africa varyingintheir habitatfrom the taxonomyofthisimportanttropical family. and theonepresented here isanothersteptowards clarifying been published( Versteegh andSosef2007 ; Recently afewtaxonomicrevisions of African generahave Couvreur 2009 ) edge of African ispoor( Couvreur etal.2006 ). and Diels(1901) andtodaythestateofourtaxonomicknowl- genera were lasttreated asawholeover100yragoby Engler Neotropics (ca.900)(Richardson etal.inpress). The African 390) whencompared to Southeast Asia (ca.1,000)andthe is theleastdiverseregion intermsofspeciesnumbers(ca. 2006 ; Punyasena etal.2008 ). Africa (includingMadagascar) rain forests around theworld( Gentry 1993 ; lianas contributingsignificantlytothediversityoftrees in Tchouto etal. observations. examination of herbariumcollectionsand additionalfield revision ofthegenus material hasbeencollected.This paperpresents ataxonomic cies nameshavebeenpublished andalarge amountof plant 1930 ). However, inthesubsequent 80yrseveralnewspe- clade consistingof genus these twobeinginturnsistertotheEast African endemic (ALBC; Fig. 1 11) andwasrecovered assisterto African generareferred toasthe African long-branchclade 2004 ). Within theLBC, the largest cladesrecognized inthefamily( Richardson etal. tered withintheso-calledlong-branchclade(LBC),oneof African Annonaceae ( Couvreur etal.2008b ), ca. five,Couvreur unpublisheddata). of 14species( Couvreur 2009 )) and Isolona eral othercloselyrelated African Annonaceae generasuchas endemic. isalso thecenterofdiversityforsev- sity liesinCameroon with fourspecies,includingonenarrow from West Africa toEast Africa. Thegenus’centerofdiver- widest distributionsknownin African Annonaceae, ranging Annonaceae isapantropical familyoftrees, shrubs, and The lastrevision of In arecent molecularphylogeneticstudyfocusing on species are illustratedandallspeciesare mapped. Conservation statusassessmentsare provided foreachspecies,aswelladiagnostickeyanddetailedspeciesdescriptions.F revision provides asynthesisofpreviously publishedinformationanddiscussionsonmorphology, , phylogeny, andpalyn species which isuniquewithin Annonaceae. Thegenusiswidespread across tropical Africa occurringinrainforest, savanna,andwoodla (9out of20species( Couvreur 2009 )), Asteranthe Keywords— Abstract— H. callicarpus (2011), 36(1):pp.33–48 contains fivespeciesoflarge trees orshrubs dis- 2

1 The New York BotanicalGarden, 200thSt.andKazimiroff Blvd,Bronx, New York 10458-5126,U.S. A. Netherlands Centre forBiodiversity(sectionNHN)/BiosystematicsGroup, Wageningen University, ale Botermans, Marleen h gns genus The . Thesethree genera were inturnsistertoa fia, Annca itiuin, distribution, Annonaceae, Africa, Isolona , occurringinMadagascar, isexcludedfrom Hexalobus Hexalobus Hexalobus Rvso o te fia Gns Genus African the of Revision Hexalobus and Monodora Generaal Foulkesweg37,6703BL Wageningen, TheNetherlands andisbasedonanextensive wasnestedinalarge cladeof consistsoffivespeciescharacterizedbysixbasallyfusedpetallobesthatare transversallyfoldedinbud, datesbackto1930( Fries Uvariastrum

3 1 Author forcorrespondence ([email protected]) Mr S M Sosef, M. S. Marc . Monodora Hexalobus Cmuiaig dtr Ade Hipp Andrew Editor: Communicating Hexalobus hasoneofthe (three outof Uvariastrum UN osrain tts, ple taxonomy. pollen, status, conservation IUCN , (6out clus- 1 Lr W Chatrou, W. Lars Hexalobus , 33 genera as also independentlyevolved in isolatedspecieswithinsuch share thisfeature ( Couvreur etal.2008b ). Sympetalyhas the monotypicendemicEast African genus Heusden 1992 ). However, petaly, is an uncommoncharacterwithin Annonaceae ( van short tubewithsixcorolla lobes.Fusionofpetals,orsym- curly hairsinterlockingadjacentsepalsinbud. The sidesoftheratherfleshysepalsare denselycovered with are, however, foundin reported by van Heusden(1992) . Reduplicate-valvatesepals never reduplicate-valvate, (i.e.margins curvedoutwards), as ( Couvreur 2009 ). Inbudthesepalmargins are valvateand while in onlyduringtheearlystagesofflowerdevelopment, trast, thesepalsinboth character shared withitssistergenus The sepalsare large andenclosethepetalsuntilanthesis,a consist ofthree sepalsand sixpetals( van Heusden1992 ). cal inshapeandare never foundtobeirregularly ruptured. examination ofseveralspecimens,thebractsappeared identi- ( Chatrou andHe1999 ), doesnotseem likely. Indeed,afterthe rupturing whentheflowerdevelops,asreported in result ofasinglebractenveloping theflowerbudandthen develop oneatatime.Thepossibilityofthisfeature beingthe obs.). Fusedbractsmightappearunusualastheynormally ( Fig. 2 ) attheirbaseandthenformatube( whenyoung.Thesebractsare free orappeartobefused Fries 1930 ; pers. upper bractsare attached nearthearticulationenclosing and sometimesproduce an additionalaxillaryshoot.Thetwo vex, soon-fallingbractsdevelopinthelowerpartofstalk genera ( van Heusden1992 ). the Annonaceae, havingbeendocumentedinseveralother less branches,oronthetrunk. Caulifloryiscommonwithin part. Thesubsessileorstalkedflowersgrow onleafyorleaf- stalk isdividedbyanarticulationintoupperandalower referred toasaflowerstalkinourdescriptions.The clustered ingroups oftwoorthree, onanaxillaryshortshoot, clades withinlarge generasuchasinspecies formerlyplaced 1999 ), and , sothegenusisnowrestricted tocontinental Africa. Thepresent Hexalobus I In As forthemajorityof Annonaceae genera, theflowers Flower budsare ovoidtoalmostglobose.Upsixcon- The usually-fragrant flowersof Hexalobus Monodora Disepalum Haplostichantus 1 n hms . . Couvreur P. L. Thomas and thepetalsare fused atthebase,forminga (Annonaceae) thesepalsalmostneverenclosepetals (Jhsn 99) 1989), (Johnson Uvariastrum Asteranthe ( van Heusden1994 ), andinsmall Asteranthe Hexalobus and Fusaea , , (Couvreur pers.obs.). Isolona Uvariastrum Isolona urthermore, four ( Chatrou andHe , , are solitaryor 2 , Monodora, 3 Sanrafaelia

enclosethe nd. The ology. . Incon- Fusea and all

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 ( 4 YTMTCBTN [Volume 36 1992 ). Theunfoldedpetalshavea characteristic“wrinkled” which isauniquecharacterwithinthefamily( van Heusden 2001). ( Rainer in SYSTEMATIC BOTANY 34 ysis (>85%)andBaysianinference posteriorprobabilities (>0.95). branches received highvaluesaftermaximumparsimonybootstrapanal- African longbranchclade(modifiedfrom Couvreur etal.2008b ). All Morton &Gledhill 898 Fig. The corolla lobesare thinandtransverselyfoldedinthebud, Fig. Raimondia 2 1 . . . . The fusedbractsina flowerbudof Cladogram ofthephylogeneticrelationships withinthe ( Westra 1995 ), butnowincludedin , WAG). , Annona

under phyllus same publication( Diels 1907 ), Additionally, twonewspecieswere alsodescribedinthat Diels (1907) synonymizedthename under after havingseentheflowers of thisspeciesafewyearslater, sister genera with thecarpelsfreely arranged.Within Annonaceae, onlythe et al.2008b ). most generaofthe ALBC ( Couvreur etal.2008a ; Couvreur pollen of variable andisreferred tohere astheconnectiveshield.The which isoftenprolonged apically. Theshapeoftheapexis oblong inshape.Thetwothecaeare unitedbytheconnective times withadarkred orpurplebasalrimontheinnerside. appearance (e.g. Fig. 3A ). Theyare cream toyellow, some- The extensiverevision byFrieswasthelast ofthisgenusand ing seeminglybilobed( Fig. 4E ). tially flat,butwithin-rolling lateralmargins, thus becom- ca. 16toppedbyasessilestigmawhichisfleshyandessen- ( Chatrou etal.2000 ). Carpelnumbervariesbetweenoneto in theNeotropical/African genera pous gynoecia(termedpseudosyncarpy, foundforexample while variousothersdevelopsyncarpousfruits from apocar- cium ( Deroin 1997 ; Couvreur etal.2008b ; Couvreur 2009 ), 1988 ), whileallspeciesin ( Su andSaunders2006 ) and Saunders 2006b ; Wanging monocarpellatespeciesinclude andSaunders2006a ), Annonaceae species( Deroin 1997 ). Other generacontain- pel. Monocarpellyisthoughttobefoundinca.10%ofthe unique inthegenus,asflowershaveonlyasinglecar- era this genushedescribedtwospecies: genus year laterby de Candolle(1832) , distinguishingitfrom the monopetala a speciesthatwaslateraccommodatedin et al.2008b ). to becommonallelevengeneraofthe ALBC ( Couvreur monocarps represent theonlymorphologicalcharacterknown to irregularly ridgedand verrucose. The sessileorsubsessile ellipsoid. Thesurfaceofthemonocarpsvariesfrom smooth sile orsubsessile,indehiscentmonocarpsthatare obovoidto are monocarpellate( van Heusden1992 ). of afruiting specimen( Engler 1899 ), to Diels, whichhadbeendescribedtwoyearsearlieronthebasis Diels (1901) transferred thespecies a newspecies, nomenclatural rules. Inthissamerevision, Englerdescribed renamed 1992 ). ter namesuperfluousandhenceillegitimate( van Heusden monopetala Candolle (1832) treated thepreviously publishedname gascariensis been transferred tothegenus and As withmost Annonaceae, thegynoeciumisapocarpous The stamensare numerous, beingnarrowly obtriangularor Txnmc History— Taxonomic The fruits of In theirrevision ofthegenus, Engler and Diels(1901) Sanrafaelia H. madagascariensis Uvaria Engl.&Diels.Thelatternamewouldbesynonymized H. bussei H. senegalensis Hexalobus A. Rich.Thegenus asasynonymof (Baill.)Engl.( Couvreur 2009 ). Unfortunately, de byitscorolla lobesfusedattheirbase.Within Isolona (Vrcut 96) 1996), (Verdcourt Hexalobus by Fries (1930) inhisrevision ofthegenus. H. salicifolius isshedastetrads,acharactershared by and as Richard (1831) wasthefirsttodescribe A. DC.Thelatterspeciesnamehas are apocarpouswithuptoeightses- Monodora Mezzettia H. monopetalus Uvaria H. senegalensis nl Adtoal, Ege and Engler Additionally, Engl. Isolona H. bussei Dielsiothamnus Hexalobus Hexalobus mossambicensis ( Verdcourt andMwasumbi haveasyncarpousgynoe- andinthemonotypicgen- Uvaria huillensis ( Diels 1925 ) as Cyathocalyx Duguetia H. senegalensis Dielsand , concordant withthe Hexalobus wasestablisheda , rendering thislat- Hexalobus H. monopetalus , and and (Wn and (Wang Pseuduvaria . However, . H. megalo- Tridimeris Engl.& Annona A DC. A. : : I. mada- Uvaria Uvaria is . ,

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 Zenker3550 and onepetalremoved. E.(abaxial surface). F. Portionofthe partoftheflowerstalkabove articulation withthre 35 BOTERMANSET AL.: REVISION OFHEXALOBUS 2011] Fig. 3 . .

; C-E, Hexalobus bussei Letouzey 7301 . A. Flowering branch.B.Stamen(abaxialsurface).- ; F, Chevalier 305 . DrawingsbyHélène Lamourdedieu. Hexalobus monopetalus . C.Floweringbranch.D.Opened flower, onesepal e monocarps.Based on: A, B, Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 J, [Volume 36 SYSTEMATIC BOTANY 36 ment ofseeds.- F. Longitudinal sectionofcarpel,showing biseriatearrangementofovules.G. Fruit with onemonocarp.H.Openedmonocarp tos Zenker 2268 Fig. 4 . .

Hexalobus crispiflorus . Modifiedfrom Flore duGabon, Le Thomas (1969) Fig.14. Hexalobus salicifolius . A. Floweringbranch. B. Flowerbud.C.Receptaclebearingstamensandcarpels.D.Stamen (abaxialsurface).E.Carpel. . I.Sterile branch.J.Monocarp.Basedon: A, C-F, Le Testu 8838 ; B, Le Testu 693 G,H, Chevalier 7471 ee biseriatearrange- ; I, Le Testu 6387 ; Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 nal tetrads(except triangular orglobose,generallylarge pollengrainsintetrago- together withtwelveothergenera,allhavingboat-shaped, placed 1972) (1971, Walker intuitive assemblyofgroups ofgenera. of thesewere basedonformal analysesofthedata,buton were basedonextensive morphological studiesthoughnone More recent classificationsofthe Annonaceae (seebelow) Thomas 1994 ; Doyle andLeThomas1996 ; Chatrou etal.2000 ). due tohighlevelsofmorphologicalhomoplasy( Doyle andLe reflects thecomplexityof itsinfra-familiartaxonomymainly common tomanyothergenerawithinthe Annonaceae, and The radicaldifferences intheclassificationof other nowknownrelated African genera(e.g. all oftheseclassificationsfailedtoinclude 37 Ophrypetalum shield-like tohalf-sphericalconnectiveappendage(notsoin ing axillaryinflorescences, valvatesepals,imbricatepetals,a the having justonewhorlofthree petals. Fries (1959) recognized ing outerpetalsaslarge asorlarger thantheinner ones,or tribe Unoneaewhichgrouped alarge numberofgenera hav- tribe Xylopieae,while Hutchinson (1923) placeditinthe or lackingaltogether. Prantl (1891) placed two valvatewhorls,almostequal,ortheinnerpetalssmall that are usuallyopenduringanthesis,petalsspread outin tribe Unoneaebasedonfloralcharacters,suchasflowers BOTERMANSET AL.: REVISION OFHEXALOBUS 1862 ) placed allied todifferent genera.Bentham( Bentham andHooker 1971). Verdcourt have beenregional inscope( Robson 1960 ; Le Thomas1969 taxonomic studieshavedealtwith ; censis, rus H.lujai treats fourofthefivespeciesnowrecognized, plusone species, 2011] 19 96) eoee recovered 1996) (1994 ; len andmorphologicalcharacters, Doyle andLeThomas and which exclusivelycomprisesthesyncarpousgenera hypothesized atransitionbetweenthisgroup andagroup tions intwotofourpartsandoilcellstheendosperm.They or transverselystriateseedswithmainlylamellarrumina- ogy. Thegenerainthisgroup havelateral,biseriate,smooth ing fiveadditionalgenerabasedonfruit andseedmorphol- Koek-Noorman (1992) placed the differences betweenthemare gradual.Van Setten and would becloselyrelated tothe laminar ovules. Van Heusden (1992) indicatedthatthisgroup free carpelsand3–16lateral uni-orbiseriatesometimes with aprominent venation, shortstamens,onetonumerous as large valvatesepals, orimbricate,ratherthinpetals era, basedonacombinationofseveralfloralcharacterssuch (1992) defineda 1992 ) andfruit ( Setten and Koek-Noorman1992 ) morphology. of pollen( Walker 1971 ; between thesefourgenerawas alsohypothesizedonthebasis Walker 1972 ), flower( van Heusden supported byhighbootstrap values.Thecloserelationship latter three genera.Theserelationships howeverwere not Isolona Based onalarge family-widesurveyofpollenmorphology, Taxonomists haveclassified In thefirstcladisticanalysesof familybasedonpol- . Thefifthofthepresently recognized species, Hexalobus Monodora wasdescribed laterby Robson (1960) . Although other . Moreover, , whichiscurrently synonymizedunder Hexalobus ) andgenerallyalateralplacentation.However, -group, including fourothergenera,allhav- . Hexalobus Uvariastrum Isolona togetherwithtenothergenerainthe Hexalobus and group, includingeightothergen- Hexalobus wasrecovered assistertothe Hexalobus Cleistochlamys Hexalobus Uvariastrum assisterto Hexalobus inthe inagroup contain- indifferent tribes Hexalobus species,theyall group andthat ) a Heusden Van ). Hexalobus Hexalobus Monodora Uvariastrum H. mossambi- Hexalobus H. crispiflo- i the in Isolona with tribe and is ).

H.crispiflorus,bussei, H.mossambicensis species of while theotherspecieshave(twoto)three to16carpels. Hexalobusmossambicensis are shorterwithadiscoid (flattened)connectiveappendage. appendage, whilethoseof oblong stamenswithanelongatedandbroadened connective a hollow“chamber.” Furthermore, tures atthebase.Bybending inwards thesestructures form corolla lobesofthelatter twospecieshavethickenedstruc- is characterized byatectumconsistingofgemmae andscabrae or two ally reticulate-rugose. The surfaceofthemonocarpsin monocarps are smoothin ornamentation provides a usefuldiagnosticcharacter. The cm wide).Unlikethewidthorshapeofmonocarps, in monocarps larger than four cmwide,whilethosefound lus lius ual pollentypes. there appeared tobea considerablevariationwithinindivid- with three pollentypesinonlyfivespecies.Furthermore, tation andinfratectumstructure. Hexalobus SEM andTEM.Thelatterstudy concludedthatpollenof five generaof African Annonaceae including pollen morphologicalvariationinamonophyleticgroup of 1983 ). Couvreur etal.(2008a) undertookanin-depthstudyof Walker 1972 ; Le Thomas1980 and transmissionelectron microscopy (TEM)( ; Walker 1971 Le Thomas1981 ; ; light microscopy (LM),scanningelectron microscopy (SEM) Le Thomas minute wartsinboth rugose in of 36cm. genus, however, are foundin H.salicifolius in distinguishing lus. H.mossambicensis ary veins,whichare indistinctonthelowersideofleaves ter. Another usefulleafcharacterisprovided bythesecond- subcordate torounded andonlyrarely attenuateinthelat- leaf basesnarrowly attenuatetocuneateintheformerand H.mossambicensis to cordate butoccasionallycuneateleafbases.Similarly, always hasacuneateleafbasewhilethelatterrounded looking species example, leafbaseshapecanhelptodistinguishthesimilar- cation in eral, leafcharactersaloneare hard touseforspecies identifi- netic analysis( Couvreur etal.2008b ). strong support,byathoroughly sampled,molecular phyloge- tionship betweenthesefourgenerawaslaterconfirmed,with studied allgeneraofthe ALBC. Nevertheless,thecloserela- It shouldalsobetakenintoaccountthatnoneoftheabove Fruits— Pollen— The pollentype observedin Flowers— Txnmcly nomtv Characters— Informative Taxonomically n and H. monopetalus Furthermore, thelengthofleafbladecanbeuseful haveflowersondistinctflowerstalks,while Hexalobus H. mossambicensis Hexalobus showssignificantdiversity insize,exineornamen- H. bussei Hexalobus Fruits are particularlyusefulfordistinguishing Several palynologicalstudieshaveincludedone (fivetotencmlong).Thelargest leavesofthe Hexalobuscrispiflorus,H.bussei, H. salicifolius speciesusingdifferent techniques suchas n and are similar. Thelargest fruits are foundin whiletheyare prominent in n and H. crispiflorus , verrucose in becausetheycanvarywithinspecies.For , althoughthoseof H. monopetalus H. mossambicensis H. mossambicensis and ischaracterizedbyasinglecarpel havesubsessileflowers.Theinner H. crispiflorus, H. crispiflorus H. bussei n and H. salicifolius H. bussei (sevenand25cmlong)from H. salicifolius, H. monopetalus Hexalobus H. crispiflorus canbedistinguishedby H. bussei andcanattainalength n and reticulately ribbedto n and are smaller (

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 1. Flowerstalk8–25mmlong;insideofbasalpartinnerpetalsnot thickened orstrongly curvedinwards, large variabilitywithincertainspecies. canbehard toidentifyasaresult ofoverlapping charactersanda distinguished onthebasisofcertainflowerand/orfruit characters.Sterile combination ofmorphologicalcharacters. 1. Flowerstalk<5mmlong;insideofbasalpartinnerpetalsthickened orstrongly curvedinwards; with light-brown hairsabove,belowprominent, covered rib impressed above,glabrous orsparselytodenselycovered covered withappressed toerect, straightorcurlyhairs;mid- occasionally sparselyhairy, lowersurfaceglabrous todensely to attenuate;uppersurfaceoftenslightlyglossy, glabrous or or acuminate,sometimesslightlyemarginate, basecordate narrowly ellipticto ovateorobovate,apexobtusetoacute young. Leaves:petioledenselytosparselyhairy;leafblade black; twigsgrey to brown, sparselytodenselyhairywhen bark rough orsmooth,fissured orflaking,brown togreyish- Hexalobus [Volume 36 10 km as criticallyendangered where thethreshold AOO undercriterionBis cell sizediameterat3.2kmorlarger willnotallowanytaxontobelisted sible valuewhichisjustunder10km of occupancy(AOO).Thecellarea wasalwayssettothelargest permis- allowed thecalculationofextentoccurrence (EOO)andthearea add-in scriptprovided byIUCN(JustinMoat,RoyalBotanicGarden Kew) ferred to ArcView ver. 3.3(ESRI2002)toproduce distributionmaps.The base ( http://dps.plants.ox.ac.uk/bol ). Subsequently, datawere trans- Willis etal.2003 ). All specimendatawere entered intoaBRAHMSdata- be usedtodeterminecategoriesofthreat usingcriterionB( Schatz 2002 ; SYSTEMATIC BOTANYthat herbariumcollectionsconstituteavaluablesource ofdataandcan ing anexpeditioninGabon. material. Thesewere complementedwithseveralfieldobservationsdur- (see Acknowledgements foralistoftheherbaria)andalcohol-preserved The pollentypeof ilar intheirhabit,habitat,andgeographicaldistribution. and agranularinfratectum,whilethesespeciesare dissim- 38 granules ( Couvreur etal.2008a ; Fig. 5 pollen typeshaveaninfratectumwithcolumellaeandsmall ). H.monopetalus verrucate torugulate tectum,whilethepollentypefoundin phologically similarspecies,ischaracterizedbyanareolate- Evergreen ordeciduoustrees orshrubs upto40mtall; Criteria usedtodistinguishamongspecieswere basedonaunique In theabsenceofdetailedpopulationinformation,ithasbeenshown The present taxonomicstudyisbasedon782herbariumspecimens 3. Leafblade5–25cmlong, 1–2.5mmindiameter;<5long,connectiveappendage slightlyconvextosem 3. Leafblade15–36cmlong,petiole 2.5–4mmindiameter;stamens6.5–8long,connectiveappendage 2. Carpel1;leafapexacutetobluntlyacuminate,basenarrowly attenuatetocuneate; 2. Carpels(2-)4–7;leafapexemarginate toobtuseorrarely bluntlyacuminate,leafbasesubcordate torounded, TYPE: 2 h ool uewt akrdo upebslrmisd;mtr ooap eeal mwd . the corolla tubewithadarkred orpurplebasalriminside;mature monocarpsgenerally>4cmwide mature monocarpsoften< 4cmwide . (SadrsadPttosWrigGop 2006 ). (Standards-and-Petitions-Working-Group 4. Leafblade7–25 cmlong,2.5–8.5wide,baserounded tocordate oroccasionallycuneate;corolla lobes>35mmlong; 4. Leafblade5–10cmlong,1.5–3.5 cmwide,basecuneate;corolla lobes<30mmlong;stamensca.2carpels3–4; ooap moho ercs . monocarps smoothorverrucose rugose (Cameroon) . . elongated andbroadened, oblique,slantinguptowards theadaxialside;monocarpsirregularly ridgedandstrongly secondary veinsindistinct() . rarely attenuate;secondary veinsoftenprominent, atleastbelow(widespread throughout Africa) . . Hexalobus monopetalus A. DC.,Mém.Soc.Phys.Genève5:212.1832.— stamens 3–5mmlong; carpels7–16;monocarpssmooth; seedswithflatraphe(West andCentral Africa) . . monocarps verrucose; seedswith raisedraphe(Central Africa) . . hasapsilateperforatetectum.Thelattertwo Mtras n Methods and Materials Txnmc Treatment Taxonomic H. salicifolius 2 (celldiameter=3.16km).Settinga n and (A.Rich.)Engl.&Diels. Hexalobus H. crispiflorus speciescanbebest Key totheSpeciesof to mor- two , tropical Africa, between15°Nand25°S. an upperandlowerpart. often slightlydarkerbrown, ruminations lamellar, dividedin brown, dulltoslightly shiny, glabrous, rapheflatorraised, tened, ellipsoidtobroadly ovoid,orange-brown tomedium seeds 2–36,lateral,biseriateorrarely in1row, slightlyflat- times constrictedbetweentheseeds,rounded atbothsides; ellipsoid tocylindrical,rarely transversallyellipsoid,some- Monocarps 1–5(–8),subsessileoronashortstipe,obovoid, appearing bilobed,hairyexceptontheinnerandupperside. fleshy, essentiallyflatbutwithinrolling lateralmargins ial sideorsemiglobose;carpels1–16,denselyhairy; apical depression oroblique, slantinguptowards theadax- tive appendageoftenelongated,discoid,sometimeswithan rowly obtriangularoroblong, extrorse, glabrous, connec- flat orconcave;stamensnumerous, free, 1–8mmlong,nar- versally foldedinbud,margins crinkled;torus cylindrical, yellow, sometimes withdarkred orpurplebasalrim,trans- rounded atapexand fleshy, withprominent veins,cream to equal orslightlyunequallobesthatare narrowly lanceolate, petals connateatbase,formingashorttubewith6distinct, covered withappressed orcurly, light-colored, minutehairs; of thebracts,insidedenselysetwithminutewartswhendry, tip, fleshy, outsidecovered withindumentumsimilartothat nate atthebase,convex,broadly ovatetowithacute round withacutetips;sepals3,valvate,free orshortly con- their baseandthenformingashorttube,broadly ovateto and enclosingthebudwhenyoung,thesefree orfusedat most bractsalmostatthesamelevelneararticulation densely covered with<0.1mm,oftencurlyhairs,twoupper- long, insideglabrous tosparselyhairy, towards themargins ered withappressed, usuallyrusty-brown hairsof0.1–1mm size towards thebase,soonfalling,outsidedenselycov- usually alternating,oftenpartlyclasping,decreasing in eral, upto6,attachedthelowerpartofflowerstalk, andoftendevelopingavegetativeshoot;sev- grant; budsovoidtoalmostglobose;flowerstalkbearing leafless branches,oremerging from thetrunk, usuallyfra- two orthree, terminalonanaxillaryshortshootleafyor surfaces. Flowersbisexual,solitaryorclustered ingroups of closely reticulate, varyingfrom distinct toindistinctonboth tal veinandoftenformingadistinctloop;tertiaryvenation the midrib,curvedupwards, eachjoiningwiththenextdis- sometimes glabrous; secondaryveins5–19oneachsideof with appressed orrarely erect, brownish orwhitishhairs, Hexalobus H EXALOBUS . . hasfivespeciesandisdistributedthroughout 2 .

4 . . . . . 3 ...... iglobose;

.... . H. mossambicensis H. monopetalus H. crispiflorus H. salicifolius H. bussei

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 areolate-verrucate tectum. F. tum. PollentypeB: C,D. and detailofpsilate perforatetectum.Scalebars: A, C,G=10µm;B,D-F, H=1µm. 39 BOTERMANSET AL.: REVISION OFHEXALOBUS 2011] Fig. 5 . . Pollen typesof H. crispiflorus Hexalobus. H. crispiflorus Pollentype A: A, B. ( de Wilde 7909 ( Liben 2390 , WAG): tetrad anddetailofareolate-verrucate tectum. E. , WAG): detailofregulate tectum. PollentypeC:G,H. H. mossambicensis ( Gomes &Sousa4897 , COI):tetradand detailofgranulartogemmatetec- H. monopetalus H. crispiflorus ( ( Chevalier 13385 Breteler 7288 , WAG): tetrad , P):detailof Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 once inMayand onceinSeptember. August, September andNovember;fruiting hasbeenreported from sealevelupto200maltitude. forests, oftengrowing inprimary, riverineorgalleryforest, Cameroon, inthesurroundings ofKribiandBipindi( Fig. 6 ). the seedorslightlydarkerbrown. Figure 3 A, B. shiny, orange-brown, rapheslightlyraised,concolorous with to ovoid,23–28mmlong,17–19wide,8–10thick, brown, shorthairs;seeds7–16,biseriate,flattenedellipsoid late patterntorugose, orange,sparselycovered withrusty- 4 cmindiam.,strongly butirregularly ribbedinareticu- Monocarps 2–4,subsessile,ellipsoid,5.3–7.8cmlong,atleast ing inahorizontalposition,3–4.6mmlong,1–4wide. hairy; stigmasattachedtothetopofovary, oftenbend- oblique; carpels3–7,4.5–5mmlong,1–1.3wide,densely 1.5 mmwide,connectiveappendageelongated,broadened, long hairs;stamensnumerous, oblong,6.5–8mmlong, base seeminglyglabrous but infactcovered with<0.1mm ones, innersidecovered with appressed, ca.0.1mmlonghairs, ish, ca.0.1–0.2mmlonghairs,apicalhairsshorterthanbasal long, 8–10mmwide,outsidecovered withappressed, whit- ceolate, 3–5mminlength,rounded atapex,23–40mm long hairs;corolla paleyellow, basedarkpurple,lobeslan- wide, outsidecovered with cream tobrown, 0.1–0.5mm ca. 15–17mmindiam.;sepals16–20long,11–14 mm 0.2–0.3 mmlonghairs;budsbroadly ovoidtoalmostglobose, in diam.,denselycovered withappressed, rusty-brown, ca. articulation 6–7mmlong,4–5indiam.,fruit 6–8mm wide, otherbractssmaller;partoftheflowerstalkabove fused, ca.6–13mmlong,third bractca.5mmlong,3.5 [Volume 36 forming ashorttubeof1–2mmlong,soonfallingexceptwhen upper bractsfree orseemingly fusedattheirbaseandthen 4–5 mmindiam.;bractsseveral,darkbrown whendry, two below thearticulation4–5mmlong,2–3indiam.,fruit grant; flowerstalk10–12mmlong;thepartof or sometimesclustered in groups oftwoonshortshoots,fra- late, distinctonthelowersurface.Flowersusuallysolitary of 55–85°withthemidrib;tertiaryvenationcloselyreticu- secondary veinsdistinctoneachside,12–17pairs,atangles hairs above,belowcovered withappressed, lightbrown hairs; 0.2 mmlonghairs;midribcovered withsmall,erect, white dry, glabrous orsparselycovered withappressed, brown, ca. glossy, oftengreyish, glabrous, lowersurfacebrown when date, coriaceous;uppersurfacesometimesslightlybullate, SYSTEMATIC BOTANY 15.5–36 cmlong,5.5–10.5wide,apexacuminate,basecor- long, brownish hairs;leafbladeobovatetonarrowly obovate, wide, sparselycovered witherect orappressed, 0.1–0.2mm young. Leaves:petioleshortandthick,1–7mmlong,2.5–4 to grey-black; twigspalebrown todarkbrown, hairywhen bark smooth,fibrous, peelingofinnarrow flakes,grey-brown Hexalobusmegalophyllus Hexalobusbussei 40 Phenology— Ecology— and Habitat Distribution— Evergreen tree of20–30mtall;trunk slender, dbhunknown; WRSL!, WU,Z!). BR!, COI!,F!,G!,HBG!,K!,M!,MO!,P!,S!,US!,WAG!, Bipindihof”, 1904, 1907. —TYPE:CAMEROON.“Kamerun: LokundjeThal, 1904 (datafrom protologue), CAMEROON. “UrwaldamUferdesKribi-Flusses”,Sept Flowering isreported from March toJune,in Only knownfrom theSouthProvince, Diels,Bot.Jahrb.Syst.39:479.1907.—TYPE: Engl.&Diels,Bot.Jahrb.Syst.39:479. Zenker 2889 A speciesfrom tropical lowland rain Busse 3216 (holotype:B!;isotypes: (holotype:B!). Zenkers.n. Zenkers.n. area ofoccupancy(ca.60 km baria representing lessthanfivelocations,andhasasmall Hexalobusbussei Hexalobusgrandiflorus Hexalobuscrispiflorus (US); Mbiave,nearBipindi,Nov1907, Zenker4831 COI, G,HBG,M,MO,P, S,US,Z);1908, river, Nyabessan,19 Apr 1968, (BR, HBG,K,LISC,M,MO,P, WAG); 30kmESEofKribi,N Kienke (BR, P, WAG); 16kmfrom Kribi,NofLolodorfroad, 19Sep1969, Kribi, NofLolodorfroad, SFIA forest exploitation,7 Aug 1969, Kienke river, justEofKribi,12May1969, smooth monocarps. and strongly rugose, theotherspecieshavingverrucose or mentation ofthemonocarpsistypicallyirregularly ridged the longerstamens(6.5–8mmlong).Furthermore, theorna- the genusbyitslarge leaf blade(15–36cmlong)aswellby Therefore the“endangered” categoryseemsjustified. is likelytohavehadanegativeeffect onitspopulationsize. estation withintheregion ( van Gemerden etal.1999 ), which for agriculture isleadingto forest degradationand/ordefor- tree). Inaddition,uncontrolled loggingandlandconversion much lessabundantnoworjusthard tocollect(alarge-sized van Gemerden etal.1999 ). Thisimpliesthat region ofCameroon tookplaceregularly afterthatdate(e.g. back to1969.Botanicalcollectingefforts inthesouth-western rence (ca.530km IC Cnevto Status— Conservation IUCN Hexalobusbussei Uses— names— Vernacular Fig. SpecimensExamined— 468 andFig.49. 1862(note:thedescriptionof thisfigure isotypes: P (4sheets)!,G!). D’hiallon [Djallon]”, Apr 1838, Cuba, Bot.Pl.vasc.1:43.1845.—TYPE: .“Fouta 6 . . (S); Apr 1908, (US);Bipindi, Apr 1910, oe recorded. None Dsrbto o of Distribution (BR,G,HBG,P, S);Mimfia,nearBipindi,Jun1899, isonlyrepresented by15collectionsinher- canbedistinguishedfrom otherspeciesof 2 ). Thelastcollectionofthisspeciesdates Zenker s.n. Benth.,Trans. Linn.Soc. London23: oe recorded. None Hexalobus bussei CAMEROON. SouthProvince: Sbankof A. Rich.,inSagra, Hist. phys. Letouzey 9387 Zenker s.n. (US);Mimfia,nearBipindi,Mar1910, 2 ) andasmallarea ofoccur- Endangered: B2B1ab(iii,iv). Zenker s.n. Bos 4495 Heudelot 865 . . Zenker 3592 (P);1908, (US). (US);Bipindi, Apr 1907, (P, WAG); 20kmfrom H. bussei (G,K,US);1913, Zenker 3550 (holotype:P!; Zenker s.n. s either is Bos 5157 Bos 5370 (BR,

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 long, 1–1.3mm wide,denselyhairy;stigmas attachedtothe age semiglobose toslightlyconvex;carpels 7–16,2–4.5mm apex, 3–5.1mmlong,0.5–0.8 mmwide,connectiveappend- numerous, oblongorsometimesslightlybroadening towards sparsely covered withcurly, fine,white, longhairs;stamens ered withappressed, ca. 0.1mmlonghairs,baseglabrous or long hairs,apicalhairsshorter thanbasalones;insidecov- outside withappressed, lightcolored tobrown, 0.1–1mm rower, 37–60(–80)mmlong,6–21wide,rounded atapex, equal orsometimessubequalwiththoseoftheinnerwhorlnar- tube 4–10mmlong,lobesnarrowly tobroadly lanceolate,all curly hairs;corolla cream toyellow, basalrimdarkpurple, (0.5–)0.8–1.5(–2) mmlonghairs,insidedenselycovered with with <0.1mm,appressed, rusty-brown hairsandwithlarger, (–16) mmwide,reflexed atanthesis,outsidedenselycovered more orlessglobose;sepals12–16(–21)mmlong,(5.5–)9–12 golden torusty-brown, ca.0.5mmlonghairs;budsovoidto fruit 4–10mmindiam.,denselycovered withappressed, above thearticulation4–17mmlong,1.2–3indiam., in fused, 8–12mmlong,4–9wide;partoftheflowerstalk then formingatubeto4mmlong,soonfallingexceptwhen 5–6, twoupperonesbarely todistinctlyfusedatbaseand mm long,1–2indiam.,fruit 4–9mmindiam.;bracts long; thepartofflowerstalkbelowarticulation2–13 or branchedshortshoots,fragrant;flowerstalk12–25mm of 42–75°withthemidrib.Flowers1–3togetheronsimple secondary veinsdistinctoneachside,9–19pairs,atangles hairs above,belowcovered withappressed, lightbrown hairs; of ca.0.3mmlong;midribcovered withsmall,erect, white when dry, sparselycovered withappressed, lightbrown hairs bullate, glossy, oftengreyish, glabrous, lowersurfacebrown times cuneate,coriaceous;uppersurfacesometimesslightly acuminate withablunttip,basecordate torounded orsome- tic toovateorobovate,7.2–25cmlong,2.5–8.5wide,apex in diam.,covered withbrown towhitishhairs;leafbladeellip- hairy whenyoung.Leaves:petiole2–8mmlong,1–2.5 ing orange-brown whencut; twigs,leavesandpetiolesdensely row, longpaperyflakes,brown torusty-brown, quicklyturn- bark ca.5mmthick,longitudinallyfissured, peelingoff innar- channelled (givingittheappearanceofabundlecolumns); Hexalobusmbula Hexalobuscrispiflorus Hexalobusgrandiflorus 01 BTRASE L:RVSO FHXLBS 41 Hexalobuslujae BOTERMANSET AL.: REVISION OFHEXALOBUS 2011] Evergreen tree upto40mtall;trunk upto1 m dbh,deeply isolectotypes: COI!,LISC(5sheets)!,LISU). 1917, TYPE: . “BucoZau,Fazenda Alsyra”, 20Jan date, Berg. 10:71.1930.—TYPE:CAMEROON.Nolocation,no BR (2sheets)!). nated: K!;isotypes:GH!,P (2sheets)!). “Ambas Bay”,1861, onymised under “Sankuru”, Jun1910, 1914.—TYPE: DEMOCRATIC REPUBLICOFCONGO. WRSL). Aug 1899, DEMOCRATIC REPUBLICOFCONGO.“Bonga,Sanga”, Engl. Monogr. afrik.Pflanzen-Fam.6:57.1901.—TYPE: says errouneously “ Deistel 99 Gossweiler 6939 DeWild., Bull.Jard. Bot.ÉtatBruxelles 4:389. Schlechter 12685 Exell,J.Bot.70,suppl.Polypet.:206.1932.— (holotype:B!;isotype:GH!,M!,P!). subsp. H. monopetalus Benth.var. H. senegalensis (lectotype,here designated:BM!; Luja, s.n. Mann 709 strigulosus (holotype:B!;isotypes:BR!, inaequilaterifolius , (holotype:BR!;isotypes: .TP: CAMEROON. ).—TYPE: (lectotype,here desig- ” anamewhichissyn- R.E.Fr., Acta Horti Engl.in (Yaoundé) ( ( Souza 1988 ). Cameroon: Ow(dial. bulu)( : Akpado Akpara Lapawe(dial.yoruba ornago) is applied. Ibaji-Ojoko ()).Therefore the“leastconcern”category (Benin), JimiraandTano Ofin(), Aponmu, Akure and Republic oftheCongo))andreserves (Ohosu,Pénéssoulou Odzala (RepublicoftheCongo),laGaramba(Democratic (), Manovo-GoundaStFloris(), (),DoudouMountains,LoangoandLopéReserve several protected areas suchasnationalparks(laMarahoue collected numerous times. Moreover, ithasbeencollectedin crispiflorus reported throughout theyear. May, Julyand August, occasionallyinNovember;fruiting is Cameroon eastward, mostfloweringoccursinJanuaryto ing isreported from January, andMarch to August. From second floweringseasonfrom OctobertoDecember;fruit- from JanuarytoMarch, sometimesto April orMay, witha from sealevelto1,000m. nas, oftennearwaterorintemporarilyinundatedforests, and galleryforests, sometimesgrowing inwoodland savan- and tothesouthofDemocraticRepublicCongo( Fig. 7 ). and southtothenorthernprovince “LundaNorte”in Angola A-H. 4 Figure brown, dulltoslightlyshiny, rapheflat,slightlydarkerbrown. long, 17–20mmwide,5–9thick,rusty-brown todark tened ellipsoidwithconvexorplano-convexsides,28–40mm rusty-brown, ca.0.1mmhairs;seeds12–36,biseriate,flat- diam., smooth,glabrous ordenselycovered withvelutinous, sessile, ellipsoidtooblong,(4.2–)5–9.5cmlong,3.5–6.5in 3.5 mmlong,(1.1–)2.2–3.1wide.Monocarps1–5(–8),sub- top oftheovary, oftenbendinginahorizontalposition,2.1– 5652 Vraua Names— Vernacular Status— Conservation IUCN Phenology— Ecology— and Habitat Distribution— Fig. ); Pota(dial.bambindjere) ( 7 . . Dsrbto o of Distribution iswidelydistributedacross Africa, andhasbeen Letouzey 4433 In West Africa themainfloweringseasonis From GuineatothesouthernpartofSudan, Hexalobus crispiflorus noa Mbl ( M’bula Angola: ); Owoe(SouthProvince) ( A speciesfrom tropical rainforests es Cnen Concern. Least D.J. Harris,J.MFay516 . . Ndoum 129 Gossweiler Hexalobus Mildbraed 6939). ) Owé ); Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 ( ( Le Thomas1969 ; WilksBovonda (dial.benga)( etal.2000 Wilks etal.2000 ). Ghana:Endwa-Gvada ). Gabon:Owui(Fang) 230 Saeger 63,121 1871, 2131,2133 128 lo Libande(dial.turumbu) ( ( 1951 ); Kpalaka(dial.Zande)( (dial. kiti)( ( 4750 13425 1460016441 ( ( (dial. turumbu) ( ( bo Liwe(dial.turumbu) ( 321 Bombai boFufouwLibande(dial.turumbu) ( ( Boutique 1951 ); Bolingola(EquateurBoende)( den Brande498 den Brande498 Democratic RepublicoftheCongo:Babua(dial.lesombo)( ( Mossome iss( Musumi (dial.lissongo)( The firstspecimen, larger leavesandlongeroblongstamens. guished byitsrugose monocarpsurfaceaswellitsmuch resembles CAMEROON. East Province: Across theSanghariver from Ndakangorilla 7278 Kilibo Forest, road downtogallery forest, 12May2000, the Luanariver, 14 Aug 1965, Lunda, road crossing neartheLuanaand Andrada river. Onthe edgeof [Volume 36 SYSTEMATIC BOTANY linda) ( 558 8831518 42 1862, 2242 have chosenthefloweringspecimen asthelectotype. near theNigerriver( Cameroon, hasflowerswhilethesecondspecimencollected face ofthosein smooth monocarpsurface,incontrasttotheverrucose sur- larger numberofcarpels(sevento16)andespeciallybyits florus species, inagreement with Le Thomas(1969) . et al.2000 ). However, weconsiderthemhere astwodistinct tus ofbothspecies(e.g.considered astwosubspecies, Wilks This hasleadtosomedifferent viewsonthetaxonomicsta- similar floralmorphologyandoccurringinhabitats. folius Walker and Sillans 1961 ; Burkill 1985 ). and venereal diseases.The fruit pulp isedible( Raponda- used asfibers,antipyretic andappliedagainstskinproblems paddles, tiles,chairlegs,andgun-butts.Thebarkhasbeen 8706 Lomo (dial.kakwa)( ( Vigne 2697 Germain 584 Nannan 565 Germain 4755 Dewèvre 860 Donis 1714,2390 Aubréville 56 Rowland s.n. Hexalobusgrandiflorus Hexalobuscrispiflorus Uses— Representative SpecimensExamined— ); Munzingo(dial.babua)( ); M’bulaNdombe(dial.luki)( ); Bonfuampimbo(dial.bangala)( (WAG); Ouémé:Eguélou,16Mar2001, ); Kakwakwa( ); Praka(dial.zanda)( , bothbeinglarge trees withdeeplychannelledtrunks, canbedistinguishedbyitsgenerallylarger leafblades, Tisserant 352 The woodhasbeenusedtomakeknife-handles, ). Nigeria:Mumfo( H. bussei De Graer246 ). IvoryCoast:Chélébé( ); Kondolio(dial. Azande) ( ); Likanga(dial.bangala)( ); MukobakobaMufike(dial.tshiluba)( ); InaoloaBombaiboFufow(dial.turumbu) ). SierraLeone:Moigbwami( ); Pota(dial.bambindjere) ( ). Central African Republic:Ewoyagba(dial. Guigonis 3125 ); InaoloaBoniningo(dial.turumbu) ( ); Bofuafua-nu-Ebaré (dial.eala)( ; ; H. salicifolius ) ( Boutique 1951 ); Mpalaka(dial. Azande) ( Gérard 2635 Louis 7195 ); InaoloaLikungu(dial.turumbu) ( ; ; . Thelatter can,however, beeasilydistin- van derMeiren 27 Madoux 274 ); Kore (dial.suma)( Barter s.n. Mann 709 Turner 78 ismorphologicallycloseto ); Kpakasa(dial.Doruma) ( Boutique wasdescribedfrom twosyntypes. Santos, dos1638 Tisserant, Equipe33,1048,1437,s.n. Andrews 1418 ); Mossoumemeko(dial.issongo) ); Iloumbe(EquateurBomongo) ); Bambu(Orientale Aketi); ( . To alesserextent, Germain 8761 Troupin 1 Taylor Evrard 2144 du Bois36 ) (Bas-Congo Lukula);Etutu ) Lota(dial.kakwa)( , collectedin Ambas Bay, ) isafruiting specimen.We ); Odiobau(LagosState) Toussaint 252 ); Kepi(OrientalePoko) (COI).BENIN.Borgou: Toui- ); Kwarka(dial.azande) Robyns 597 NOA Lna Norte: Lunda ANGOLA. Goossens 3022 Aubréville 810,1794, De Wulf 705 Akoegninou 4406 ). ). EquatorialGuinea: Fay 5331 ); M’bula( Harris 1814,2451 ); BulaN’Dombe Dawe 475 Hexalobus crispi- H. crispiflorus Evrard 3203 van derMaesen ) Boniningo ); Baland 1891 ; ; ) Mosome ); Wagemans Gutzwiller Dechamps ) Kpakas ); ) : ). H. salici- ) Loopa ); Germain Nannan (WAG). Myers Louis van van de ); ). ); )

KerstingA594 Hexalobusmonopetalus Uvariahuillensis 1978, Mono river, Mar1987, 314 8706 experimental farmnear Yei, 20Mar1939, study area, 8Mar1988, Hexalobusmonopetalus Hexalobusglabrescens Hexalobustomentosus Hexalobusmonopetalus Kosper, P. 235 24 May1939, Jan 1967, Mar 1964, .about5milesfrom Alikalia onroad from Bendugu,22 National Park,bankofLékoliriver, 8Feb1994, (FHO); 1948, from Ado-Awaiye ontheIseyinRoad,28May1957, NIGERIA. Abo, SNigeria,March 1858, river withtheroad Taï-Tabou, 12Mar1962, 621 along theroad from Sankadiakro toManzanonan,1 Aug 1969, LISC, M,WAG). IVORY COAST. Abengourou: 25kmNEof Abengourou, Bafata, MadinadeMamdu Alfa, 1May1956, Cosselinta, 26May1953, 55193 WAG). GUINEA Kankan:5kmNWofKankancity, 23Dec1966, Region: betweenWenchi andBamboi,8Mar1995, the road toMvadi,5Nov2005, (K, MO,WAG). GABON.Ogooué-Ivindo:Bélinga,Iron mines,6kmon 19–20 kmfrom the Alegre river, 32NMK6207,2 Apr 1990, 3596 ings closetoBusanga,ontheKolwezi-Wakipinji track,9 Aug 1987, valley Kinkoko,11 Feb1949, (K, MO).DEMOCRATIC REPUBLIC OFTHECONGO.Bas-Congo:Luki, Sangha riverfrom NdakanCampupstream, 9Feb1989, 3125 Langandou [probably Bagandou],Gouligouliriver, 20 Aug 1964, Fay 558 (K);KhorsKagelu,16Jul1931, (MO,WAG); Tabou: LeftbankofHanariver, nearthecrossing ofthis (A);nodate, (BR).EQUATORIAL GUINEA.Bioco(FernandoPoo):Malabo-Luba, (BR,K,P);SanghaEconomique:45kmSofLidjombo,Eastbank isolectotype: BM!). no date, 6: 56.1901.—TYPE: ANGOLA. “Benguella,Huilla”, Engl. &DielsinEngl.,Monogr. Afrik. Pflanzen-Fam. Berlin. 2:296.1899.— G00011597]!, here designated;isolectotype:P!). prope Bakel”,1828, ous name—TYPE:.“Sénégambie:Galam DC., Mém.Soc.Phys.Genève5:213.1832,superflu- nated: K!;isolectotypes:BM!, Z!). French Ubangi”,1911, 52. March 1927.—TYPE:CAMEROON.“Lere toHam, designated: P!). Sansanding”, 29Sept1899, pl. 1:10.1920.—TYPE:.“Segou,PaysMandé, types: G!,P!,WAG!). M’bra”, 30Nov1902, CENTRAL AFRICAN REPUBLIC:“Environs deKaga Chiefs andcitiesofCentral Africa: 301.1912.—TYPE: Seneg. tent.:8, Fig. 2 . 1831.— 1901.— Monogr. afrik.Pflanzen-Fam.6:56,t.20,fig.Ba-d. 4536 District, E.ofMatonchi Farm”,12Feb1938, Bot. Gard. 8,3:214.1953.—TYPE:.“Mwinilunga ; Kindia,Mar1905, Ern 3231 (K,MO).CENTRAL AFRICAN REPUBLIC.RoadMbaïki- Gledhill 568 Morton SL 989 (holotype:K!;isotypes: BM!,BR!,PRE). (K);Equatoriaprovince. KageluStation,3Mar1938, Russell FHI22551 (FHO). Andrews 1418 Uvaria monopetala (P);1906, Antunes 266 Neilson 25 Engl.&Diels,Notizbl.Königl.Bot.Gart. (BR,FHO,K,WAG). SUDAN.Iszuriver:MeidiDistr, Brunel, 2109 Fay 8299 (K,WAG); NorthernProvince: BumbanNP, 22 Hutch.&Dalziel,Fl.W. trop. Afr. 1: A.Chev., Expl.bot. Afr. occ.Énum. Chevalier 13385 Espirito Santo3185 var. Kersting A392 var. Donis 2390 (K);SouthwestEquatorialprovince. Kagelu (FHO);Lado, Yei river, 23Oct1919, Leprieur s.n. (K).REPUBLICOFTHECONGO.Odzala (lectotype,here designated:COI!; Hexalobus huillensis Sosef 2225 (A.Rich.)Engl.&DielsinEngl., obovatus Chevalier 6486 Talbot 531 (F, MO,NY, P);30 Apr 1988, parvifolius A. Rich.inGuill.&Perr., Fl. Turner 78 (BR);Sotouboua, Anié river, 28Feb Chevalier 2542 (BR,K);Katanga(Shaba):surround- Barter s.n. (HBG);Tihabola Kuanda,1906, (P).GUINEA-BISSAU.Chitole, (WAG). GHANA.Brong-Ahafo Brenan, Mem.New York Hexalobus senegalensis Hoyle 789 de Wilde 3605 (FHO,LISC,WAG); Bafatá: (lectotype:G[barcode: (K). Kri-Kri,bankof (lectotype,here desig- Baker. f.inMacleod, Dowsett-Lemaire 1631 Espirito Santo3841 (K);OyoState:ca.2miles (holotype:K!;iso- Jongkind 2109 (FHO);3Feb1983, (lectotype,here (Engl.&Diels) Onochie FHI31206 Harris &Fay1814 (BR,GH,WAG). Milne-Redhead Carvalho (BR,MO, Versteegh Harris & Guigonis , , Lisowski Schaijes (FHO, de 4311 Sillitoe Myers (BR). A.

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 during theyear. Fruiting hasbeenreported inJanuary, from dant withthepercentage ofcollectionsthat havebeenmade from OctobertoFebruary. Thispatternis,however, concor- the yearwithmajorityof floweringspecimensgathered andNigeria)flowering hasbeenreported throughout the appearanceofnewleaves. InWest Africa (from Senegalto rocky places,from sea levelto1,600maltitude. nas orgalleryforests, oftengrowing onsandysoilindry absent from theevergreen forest regions ( Fig. 8 ). to thenorthofSouth Africa andthesouthwest of Angola, but brown, rapheraised. Figure 3 C-F. ovoid, 10–15mmlong,7–10wide,5–8thick,dull, biseriate orsometimesin1row, slightlyflattenedellipsoidto covered withshorthairs,fruit pulpwhitish;seeds2–8(–11), let, withminutewarts,sometimesreticulate-rugose, sparsely between theseeds,2.2–4.6cmlong,1.3–2.2indiam.,scar- cylindrical, ortransverselyellipsoid,sometimesconstricted 1–5, sessileorwithastipeuptoca.3mmlong,ellipsoid lobes withtheedgesslightlycoiledinwards. Monocarps or rusty-brown hairs;stigmas1–1.3mm long,dividedintwo 2.1–3 mmlong,0.8–1wide,denselycovered withgolden mm wide,connectiveappendagediscoid;carpels(2–)4–7, numerous, narrowly obtriangular, 1.4–2.1mmlong,0.5–0.8 with appressed, hairs,<0.1 mmlong,baseglabrous; stamens appressed, rusty-brown hairsca.0.2mmlong,insidecovered appressed, whitishhairs, <0.1mmlong,basecovered with gins reflexed toformahollow chamber, outsidecovered with lobes curvinginwards at thebaseabovetubeandmar- wide, rounded atapex, inner whorloftennarrower with lobes narrowly lanceolate, (9–)12–22(–27)mmlong,3–7 mm wide;corolla cream topaleyellow, tube2.5–4mmlong, almost globose,3–5mmindiam.;sepals4–7long,3–5.5 golden orrusty-brown hairsca.0.2mm long;budsovoidto in diam.,flowerdenselycovered withappressed, cream to flower stalkfrom hardly discernableto2mmlongand 5–6, closelyspaced,largest onesca.5mmlong,3.5wide; sometimes emerging from thetrunk, usuallyfragrant;bracts Flowers 1–3togetheronleaflesstwigsoramongtheleaves, 43 venation varyingfrom distinct toindistinctonbothsurfaces. side, 6–14pairs,atanglesof50–80°withmidrib;tertiary ondary veinsraisedanddistincttoratherobscure oneach brous orwithappressed toerect, whitetobrown hairs;sec- erect, oftencurlywhitetolight-brown hairsabove, below gla- long hairs;midribglabrous orcovered withdense tosparse appressed toerect orcurly, whitetolight-brown, 0.2–0.8mm 0.3 mmlong,lowersurfaceglabrous todenselycovered with glabrous orsparselycovered witherect cream hairsca. subcordate, coriaceous,uppersurfaceoftenslightlyglossy, sometimes lightlyemarginate, baserounded tocuneate or wide, apexobtusetorounded orrarely slightlyacuminate, BOTERMANSET AL.: REVISION OFHEXALOBUS tic toslightlyovateorobovate,3.6–17.5cmlong,1.2–6.5 usually lightcolored hairs;leafbladenarrowly elliptic orellip- ered withdensetosparseerect, oftencurly0.2–0.3 mmlong, young. Leaves:petiole1–4mmlong,1–2indiam.,cov- or orangetobrown; twigsgrey tobrown, oftenhairywhen sometimes brown oryellowish;slashfibrous, pale yellow bark rough orsmooth,fissured orflaking,generally grey, times prostrate, oftenseveralstemmed,dbhupto35cm; erect orspreading habitandlowbrancheswhich are some- 2011] Phenology— Ecology— and Habitat Distribution— Deciduous shrub orsmalltree upto10(–15)mtallwith Flowering takesplacebefore, ortogetherwith From West Africa toSudan,from A speciesfrom (woodland)savan- (Parraka) ( Tanambakada Ayetinbaga (dial.waama).:Nyei Akok fon); MwesweTibaka Ablogbe (dial.bariba)( Souza 1988 ); (dial. mundimba)( Mutundo (dial.cuanhama)( ( (dial. kitabwa)( 847 kitabwa) ( (Katanga (Shaba)Bukama)( Kalendasi-kisofu (dial.kibemba)( Boutique 1951 ); Katwelubala ( “least concern”categoryissuggestedhere. (Mutaro Kunda(Gambia), Kafarati(Nigeria)).Therefore, the (), Kruger ()) andforest reserves (Central African Republic),Comoé(IvoryCoast),Matopos (Democratic RepublicofCongo),Manovo-Gounda-StFloris as gamereserves (Yankari, (Nigeria)),nationalparks(Upemba Moreover, ithas beencollectedinseveralprotected areas such number ofherbariumspecimenshavebeenrecorded. monopetalus from OctobertoDecember. fruiting hasbeenreported from JanuarytoMarch, May, and flowering hasmainlybeenreported from MaytoNovember; , Zambia,Zimbabwe,Mozambique,andSouth Africa) In southwesternandsoutheastern Africa (Angola,Tanzania, October toDecember;fruiting isreported inJulyand August. Republic) floweringspecimensare collectedmostlyfrom Africa (Chad,Sudan,,Cameroon andCentral African March to August andinOctober. InthenorthofCentral ( 1085 Nêfê (dial.dioula)( (Upper riverDivisionFatoto) ( Da-afi (UpperriverDivision Fatoto)( Moko (dial.Mudianshiakiluba) ( de Ganays.n. Teixeira 5584 Dubois 3 Vraua Names— Vernacular Status— Conservation IUCN 8. Fig. ); Kizofwe(dial.kitabwa)( Boutique 1951 ); Kundukundu ); Fokania(Sikasso)( ); Foukania (Sikasso)( Dsrbto o of Distribution A. Schmitz959 Turner 94 iswidelydistributedacross Africa, andalarge ); Muhongu(dial.ganguela)( ); Goño(Mopti)( Schmitz 1376 Santos, dos2792 Ambe 229 ). DemocraticRepublicoftheCongo: Hexalobus monopetalus Angola: Mubebe (dial.muquipunge) Ellenberger 500 ); Kizofu (dial.kibemba)( Azancot deMenezes1312 ); Moko(dial. kisa)( Vanden Brande49 ). Mali:Fin(Sikasso)( Ellenberger 763 Rosevear 129 es Cnen Concern. Least Griaule s.n. ). Benin:Dankokwe(dial. de Troyer 126 ); Fougandier(Kayes) Rosevear 107 . . ); Gaño(Mopti) ). IvoryCoast: ); Kikobo(dial. ); Hablé(dial. Mendes 3385 Homblé 1281 ); Mutundu Mutundu ); ) Gambia: ). Ellenberger Hexalobus ) Suma ); Schmitz ); ); Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 (Woulli BaolSaloum)( Nigeria: Boileh(dial.fufulde)( Ushizou Nyiha(Mbeya)( ( duigue) ( lector number or evencollectingdatewaslacking from this assigned andspecific informationsuchasa collectororcol- French botanist Richard (1831) . A typespecimenwasnot logically distinctfrom therest. southern Angola ( Fig. 8 ) butthesespecimens are notmorpho- Hexalobusmonopetalus tribution reflects thedistributionofthesexericecosystems. ecosystems suchassavannasandwoodlands,itsdis- [Volume 36 SYSTEMATIC BOTANY ( goudibelou) ( 44 (dial. chivenda)( (Kitwe District)( Mukundu-kwibila (Ndola)( Mu-bnundu-hui-bwila (CopperbeltProvince) ( Redhead 557 madi) ( (dial. nyam)( 81 madi) ( Sudan: Lûto(dial.pari)( Shakama plumShakamapruim ( Coates Palgraveetal.1983 ). South Africa: Mulanzakamelo(dial.venda)( Akoègninou etal.2006 ). pulp isedible( Burkill 1985 ; Ruffo etal.2002 ; ing andrashes,bronchitis, andasahorsemedicine.Thefruit Arbonnier 2004 ; are usedasanexpectorant, incaseofbloodyvomiting,itch- chew sticks,buildingpoles,charcoal andfirewood. Leaves legs, toolhandles,bows,gunstocks,spoons,carvings,andas and toothdecay. Wood hasbeenreported tobeusedforchair lines. Stemsare usedagainst colds,cataracts,puerperalfever venereal diseases.Barkfibers are usedasrope andfishing antipyretic andincaseof insomnia,colic,constipationand case ofheadachesanddiabetes,asadiuretic andlaxative,an snake bites.Otherreports mentiontheuseofthisspeciesin treat stomachpainsaswell astostoppoisoningcausedby (Tanzania, Mbeya region) theroots are boiledanddrunk to the ‘Ipota’area between theMsheweandMuvwavillages contrast to aethiopica African Annonaceae. Onlyafewotherspeciessuch as more precise way. collection datamighthelptocharacterizethisvariationin a recognize anyinfraspecificvarieties.Furtherfieldworkand continuous variationbetweenforms.We thusdecidednotto lus the midribwhichisnevertomentellousasinvar. Africa, onthebasisofitsdistinctbroadly obovateleavesand var. foliage andindumentum. Brenan (1953) , however, described and itsprominent leafvenation, especiallybelow. former isdistinguishablebyitsmulticarpellategynoecium ers andsmallmonocarpswithsmoothsurfaces.However, the being smalldeciduoustrees orshrubs withsubsessileflow- Abeid 1195, 1338 Duvall 138 The protologue of Uses— ). Tanzania: Imuli(dial.kisafwa)( Hexalobusmonopetalus Hexalobusmonopetalus Hexalobusmonopetalus . Basedonthespecimensstudied, there appearstobea obovatus Brasnett 330 Jackson 520 havesuchanextensivedistribution.However, in The labelson Trochain 1390 ). Mozambique:Linkopila(Niassa)( ); Karendesa (dial.chilunda)( Xylopia aethiopica , predominantly dispersedintropical South Leclercq-Rogeon 301 Wigg 1120, 1185 Davies 1320 ); Mkuwa(dial.nyamwezi)( Holmes 688 ); Tloloyot(TheBoyahillKimatong)( ). Zambia:Kanpo(dial.chikaonde)( Uvaria monopetala alsooccursinan isolatedregion in ); Poili(dial.pench)( hasoneofthewidestdistributionsfor Lovett 4040 isaspecieswhichvariableinits Heudelot 360 resembles Simpson 7512 Knight 76 ). Fanshawe 920,1299 , , ). Zimbabwe:Mutanzakameru H. monopetalus Dalziel 205 ); Mkutowa(dial.kitongwe) ); Kungé(dial.malinka) & Lovett 4040,4317 H. mossambicensis ). Uganda:Erota (dial. ); Coundié(dial.man- waspublishedby the 4317 ); PopiReta(dial.ku Milne-Redhead 4536 ). Senegal:Casséass mentionsthatin grows indrier Trochain 1158 Gerstner 5911 ) Mu-lnundu ); Holmes 658 Jansen 7891 Msese 4846 monopeta- ) Mkua ); Xylopia Milne- both , Fukui ). ); ); ); ); ). as wellthename tained localityinformation:“Galamprope Bakel”(Senegal), given acollectornumber. Severalofthesespecimenscon- botanist Leprieur. Unfortunately, thesespecimenswere not imens were collectedinSenegal1828byanotherFrench ties: nearGalamandJoal(Senegal). A numberofspec- protologue. Richard (1831) , however, mentionedtwolocali- the protologue of and Diels(1901) , onlythelocalityinformation,copiedfrom by De Candolle(1832) orof quent publicationsoftheprotologue of several ofthesespecimenscollectedbyLeprieur. Insubse- it seemslikelythatRichard basedhisdescriptionononeor same nationalityofthecollectors,timeperiod,andlocality a specieshenamed Swiss-borne French botanistnamedPerrottet also collected totype isthattheoriginalspeciesname, Hexalobusmonopetalus collected byLeprieur(from Senegambia)from P asthetypeof Fries (1930) andlater Verdcourt (1971) mentionedaspecimen ( Lanjouw andStafleu1954 ). This Chevaliercollection,however, Types describedbyBaker f.are generallystored inBMandK 6486 described byBakerf.in1912. He citedthespecimen strong. elsewhere westillfeelourchoiceofalectotypeissufficiently rying thenameJoaland/or monthofMaymayturnup but wewere notabletolocatethis. Although aspecimencar- Verdcourt (1971) mentionedtheexistenceofanisotypeinW, moreover isagoodrepresentative ofthespecies.Inaddition, specimen sinceitwasmostlikelyobservedbyRichard and in theprotologue. We assignthestatusoflectotypetothis collected inrocky hills,informationthatwasalsopresented locality “Galam”itiswritteninoldhandwritingthatwas specimens from P orG).Onthelabelnexttocollecting branches resembles thedrawingwhencompared totheother ilar tothedrawinginprotologue (theconfigurationofthe flowers. Oneofthesefloweringspecimens,G00011597, issim- G00011596, isafruiting specimen,whiletheothertwobear monopetala lecting locality:“Galam”aswellthecorrect name G00011597) three duplicatesbearalabelwith thecorrect col- have examinedfrom G(G00011593, G00011595, G00011596, both specimensbearthename P00315832) donotmentionanycollectinglocality, although the othertwoduplicatesfrom P weobserved(P00315836, this specimencannotbegiventhestatusofholotype. Also Richard basedhisdescription andwetherefore argue that tioned. We assume thatthiswasnotaspecimenonwhich not indicated. Also thelocalitiesGalamorJoalare notmen- On theoriginallabelhowever, thename was literallycopiedfrom the protologue of type?” andmentionsthecollectinglocality, informationthat (barcode: P00315834). A recently addedlabelsays:“typeholo- herbarium ofRichard, and whichwaslatertransferred toP the collectinglocalityandyear. based hisdescriptiononit).Otherimportantinformationis to knowifRichard hasobserved thespecimen(andideally, indicated onan“original”label.Furthermore, wewouldlike The prerequisite ofaspecimenwhichcouldbecomethelec- The variety Of thefourduplicatesofLeprieurcollectionwe We haveseenacollectionbyLeprieurthatbelongedtothe butdidnotindicate theherbariumwhere it waskept. (G00011595, G00011596, G00011597). Oneofthese, Hexalobus monopetalus Uvaria monopetala Uvaria monopetala Uvaria monopetala . Hexalobus monopetalus Uvaria monopetala , isgiven.However, first . However, in1827the var. inGalam.Giventhe Hexalobus senegalensis Uvaria monopetala Uvaria monopetala Uvaria monopetala parvifolius . by Engler Chevalier Uvaria was , is , , is , .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 1968, of river13Oct1989, Gouda-St. FlorisNationalPark,1kmNofKoumbalacamponside WAG). CENTRAL AFRICAN REPUBLIC.Bamingui-Bangoran:Manoro- map #NC33VIIIGaroua, 400m,13 Aug 1983, 13 Aug 1983, 10 Sep1969, Katanga (Shaba):Lubumbashi,14Feb1950, Mara, 1903, (K); MoyenChari:Charicentral,LacIro region, from KabaSolàto 1998, Essou2360 1959, 21 Jun1949, Mitwaba area. Ganzaleftaffluent Lukokariver. UpembaNationalPark, classified forest, 21Nov 1984, 22511 about 10milessouthofKishionIgbohomotorroad, 29Feb1948, 15 Jul1973, Adamawa Province, onthe mainroad to Yola, 5milesfrom MayoBelwa, Bankangau, Mar1983, Leistner, Oliver, Steenkamp&Vorster 229 32 01 BTRASE L:RVSO FHXLBS 45 (= Vila Arriaga), nodate, BOTERMANSET AL.: REVISION OFHEXALOBUS is congruent withthehabitofthisspecimen. men istheholotype,takingintoaccountthatdescription folius in 1907. Although thename is notinBMbutthere isaspecimeninKwhichwasacquired 2011] Otijemba, 17Jun1965, 57 (M); SouthWest Africa. Swartbooisdrif1713BD,16Jun1968, south ofRuacana,14 Apr 1966, WAG). .SouthWest Afrika.1714 AC RuacanaFalls.15km District. Montepuez,MonteCherimpa,2Jan1964, 1971, 1899, river Bakhoy, 1Jan1948, 16 Oct1979, 1988, Province: Mt.MulanjeMassif.MichesiMAP REF.: 893554,4 Aug 4 milesSEofMzambazi,22Jan1978, 1978, ca. 16kmNorthofKorhogo, justaftervillageLataha.Guineazone,2Nov 10 kmNEofBouin,11 Feb1968, Piche -Buruntuma, 9Dec1961, Wa-Sawla road, 5Nov1958, MortonA 768 Region: BetweenPepeaseandNkwantanang(Kwahu),14 Apr 1954, Rosevear107 23 Feb1995, 16 Feb1949, 1954, Falls, 10Feb1935, feet elevation,Dec 1931, Collected intheRuduorsacred grove atMoyo.Madidistrict. About 3.800 1971, 1983, Kanté, mountainsWofTaberna country, 15kmNWofKouatié,16Nov Badiouré (lowerCasamance), 10m,2Jun1982, ders thenamevalid. which is,thoughminimal,acharacterofthisspeciesandren- a nomennudum,butitsprotologue states“arbre” [=tree], Muvwa villages,2Feb1990, Vollesen MRC4908 Schweinfurth2397 foothills, 12Nov1948, M). SUDAN.EasternEquatoria:Equatoria,Torit, Molongori, Imateng date, 7432 Zoutpansberg Nords. nearsaltpan,ca.1,100m,20Oct1955, Transvaal. Zoutpansberg District.52mileswestofLouisTrichardt Sine-Saloum: Kaolak,1904, (M);Kaokoveld,Ondsorusu Falls ontheKuneneriver, 22Jul1976, (M);Kaokoveld:KaokoveldKAO.Baynesberge, 3milessouthof Fnly Finally, Representative SpecimensExamined— (HBG,K,M,NY, US);Transvaal. Dist.Waterberg. Spruytskloof, no (FHO,K).SENEGAL.Tambacounda -Kedougou,km74Diambour Breteler 7288 Fotius 1085 Schäfer 7760 Theron 2063 Chevalier 274 Chapman 9251 Ellenberger 1085 Fanshawe 1559 Teixeira 4423 isnotindicatedonthelabel,weassumethatthisspeci- Dekker 320 van derMaesen6556 (WAG); Zou:13kmSofTchetti, Kitikpli,alongroad, 20Nov Chevalier 9099 Witte 6882 Chapman 5 Hexalobus tomentosus (FHO);27 Apr 1936, Bognounou 233 Fox 37 Descheres 94 Thomas 2432 van Melle48 (GC,K,WAG); NorthernRegion:2.5milesSofKulmasa, (P);Parraka-Zaude,KudoDinka,20Jan1932, (WAG). MALAWI. NorthernProvince: MzimbaDistr, (K,P).TANZANIA. Lindi:NahombaValley, 8Feb1978, (MO,WAG); Center:km37 Sokodé-Lamakara,8Nov (M);Waterberg District,Dec1956, (B,BR,MO,US,WAG). UGANDA.NorthernProvince: (P, WAG); Sikasso:4kmEdeKofoulabé,20February (BR,COI).BENIN. Atakora: Péhunco,16 Aug 2000, (K);LowerriverDivision:Mutaro KundaForest Park, (C,EA,K);Mbeya:Ipotaarea betweenMsheweand Taylor 3388 (FHO,MO,WAG). MALI.3km.WofTomanibugu, (BR,FHO,K);Central Province: Broken Hilltown, (P, WAG). MOZAMBIQUE.CaboDelgado:Niassa Harris 2046 (BR,WAG). GAMBIA.nodate, (FHO,K);OyoState:Province, OyoDistrict, Jackson 520 Boudouresque 5981 Giess 8998 (BR);UpperriverDivision:Fatoto,27 Apr 1936, (K,P).DEMOCRATIC REPUBLICOFCONGO. Brasnett 330 Roberty 10307 (MO,P, WAG). CAMEROON.NorthProvince: (K,P, WAG); from KitatoGallé,edgeofthe (P);NorthProvince: map#NC33VIIIGaroua, Antunes 32 (WAG). . Comoé:Banfora, Kaichinger 5 Lovett 4040 Rose Innes1149 (MO,UPS,US).ZAMBIA. Ndola,14Sep Bamps 7711 Pereira 2261 Giess 1714 (MO,US,WAG). CHAD.Golé,16Oct Geerling 1994 (M);1955, (FHO);Western Equatoria:14Jul1969, Hexalobus monopetalus Rosevear 129 Pawek 13665 A. Chev. isoftenregarded as (K);Western Province: Murchison (COI);Moçamedes,Lungo,28Jul (G,MO,WAG). TOGO. Reg.Kara- (G);Koulikoro: Keniegue,26Jan (P). SOUTH AFRICA. Transvaal: (P).NIGERIA. Adamawa State: (MO,US).NIGER.ParkW (BR,MO,WAG); Casamance: (K).GUINEA-BISSAU.Gabu: (M);14 Apr 1966, (C).IVORY COAST. Bouna: NOA Nmb: Bibale Namibe: ANGOLA. Koehne 33 Schmitz 2491 (C,MO,WAG); Korhogo: (FHO).GHANA.Eastern (BR,MO,WAG); Southern Vanden Berghen 5203 Thomas 2432 Torre 9831 Dawe 51 Theron 2025 (M);1955, (BR,COI,K); var. (P);Genieri, (BR,MO, Giess 9299 Grobbelaar Turner 94 (MO, P, Keay FHI Schlieben (FHO, Koehne parvi- (BR);

Davies1320 (SRGH, WAG); Beitbridge.Horseshoevalley. Limpoporiver, May1955, of theconfluenceMusirizwiandBavazirivers,29Jan1975, 13 Sep1947, and October. December, butmainlyinOctober; fruiting: Januaryto April soil, at20–200m. the dunesnearcoast.Growing onbrightorred, sandy forest, insecondarywoodlandorthewoodyvegetation of Cabo Delgado.( Fig. 9 ) uted northoftheZambeziriver, inprovinces Nampulaand darker brown. orange-brown, dulltoslightlyshiny, rapheflat,slightly semiovoid, 18–22mmlong,13–15wide,5–8thick, brous, denselysetwithminutewarts;seeds10–14,biseriate, sessile, ellipsoid,2.8–6.7mmlong,2–3.7cmwide,gla- hairy exceptinsideandontopofcupule.Monocarp1,sub- cupuliform, openononesidewiththeedgescoiledinwards, densely hairy;stigmaca.1mmlong,1.5wide,broadly nective appendagediscoid;carpelsolitary, 2–3mmlong, rowly obtriangular, 1.8–2.4mmlong,0.4wide,con- to 0.1mmlong,appressed hairstowards apex;stamensnar- lobes glabrous atbase,getting sparselycovered withshort, towards apexwithca.0.2 mmlonghairs;innersidepetal 0.5 mmlonghairsatthebase,gettingmore sparselyhairy lobes denselycovered with appressed, lightcolored, ca. base andcoveringreproductive organs, outersidepetal ened structures attheirbase,alllobes bendinginwards at inner petals,thelatterslightlynarrower with2ovoidthick- at basewhere lobesofouter petalspartlyoverlapthoseof 10–18 mmlong,2–3.5wide,rounded atapex,widened whitish hairs,insideglabrous, lobesnarrowly lanceolate, 1.5–3 mmlong,outsidesparselycovered withappressed, densely packedcurlyhairs;corolla cream toyellow, tube ca. 0.5mmlonghairs,insidesparselyhairy, margins with long, 3.5–5.5mmwide,outsidewithlightbrown towhitish, hairs; budsovoid,ca.3–4.5mmindiam.;sepals6–8 long, denselyhairywithappressed, lightbrown towhitish the partofflowerstalkabovearticulation0.5–4mm outside, insidecovered with sandycolored suberect hairs; covered withrusty-brown togreyish appressed hairsonthe eral, largest onesca.5mm long,3.5mmwide,darkbrown, flower stalkbelowthearticulation1–2mmlong;bractssev- or leaflesstwigs;flowerstalk1.5–5mmlong;thepartof tinct onbothsurfaces.Flowerssolitary, subsessile,onleafy angles of45–80°withthemidrib,tertiaryvenationindis- long; secondaryveinsindistinctoneachside,6–12pairs,at covered withappressed, light-colored hairsofca.0.2mm ate tocuneate,coriaceous,glabrous above,belowsparsely or sometimesacutewithrounded tip,basenarrowly attenu- (1.1–)1.5–3 cmwide,apexusuallyacuminatewithblunttip narrowly elliptictoovateorobovate,(3.2–)4–9.3cmlong, 2–5(–8) mmlong,1indiam.,sparselyhairy;leafblade and petiolesdenselyhairywhenyoung.Leaves:petiole branched, dbhunknown;barksmooth,greyish; twigs,leaves Hexalobusmossambicensis Phenology— Ecology— and Habitat Distribution— Deciduous shrub orsmalltree upto5(–15)mtall,much (holotype: LISC!;isotype:BM!). Fernão Velozo eQuissangulo”,15Oct1948, 2, 32:153.1958.—TYPE:MOZAMBIQUE.“Nacala,entre (MO). Brenan 7856 Flowering in April andSeptemberto Endemic toeasternMozambique,distrib- (FHO).ZIMBABWE.ChipindaDistr. HillW A speciesgrowing inclosedoropen N.Robson,Bol.Soc.Brot., sér. Barbosa 2428 Biegel 4855

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 Mocimboa dePraiatoDiaca,14 Apr 1964, 1964, Mocimboa beach.5kmfrom MocimboabeachtoDiaca,(Kb),20m,14 Apr Msala riverandMoeiuiboa,12Sept1948, EA); Niassa:6Sept1948, and thebeach,infront of the mission(Kg),25Jan1968, 17 Oct1965, 1984, de Feb 1984, Forest Reserve,20Feb1984, e Sousa4863 4897 of columns);bark ca.6mmthick,longitudinally fissured, deeply channelled, (givingittheappearance ofabundle Hexalobussalicifolius Dec 2003, GomeseSousa4670 Diaca-Chai road, 3kmof theriverMuera,onleftedge,26Mar1961, [VolumeMatibane). Eventhoughthe 36 AOO isjust120km national park(Quirimba)andonefloralreserve (Meorusse de In addition,twocollectionswere madeinprotected areas, one herbarium collections(14)includingarecent onefrom 2003. Hexalobusmossambicensis SYSTEMATIC BOTANY 46 are, however, notbilobedinshape. It shares thelattercharacter with bilobed, thickenedstructures atthe base oftheinnerpetals. flowers andleaveswithindistinctveins.Furthermore, ithas other speciesinthegenusbyitssolitary, monocarpellate used fortheconstruction offurniture ( e Sousa4670 than an“endangered” one. in twoprotected areas suggestsa“vulnerable”statusrather Evergreen tree upto35mtall;dbhtrunk upto1m, Status— Conservation IUCN Fig. Hexalobusmossambicensis Uses— Names— Vernacular SpecimensExamined— (COI); António Enes(Angoche),nearthecoast,15Oct1965, K!, M!,MO!,P!,WAG!, WU,Z!). Zenker2268 Fam. 6:57.1901.—TYPE:CAMEROON. “Bipinde”,1900, Torre 11928 9 . . Koning 9655 Groenendijk 1157 Luke 9942 The fruits are edible( Dsrbto o of Distribution (BR,COI,EA,K);MossurilDistrict.Meorusse toMatibane Mogg 32398 ); Nencopilo(Nacalaregion) ( (COI,EA); CaboDelgado:MocimboadaPraia.5kmfrom (COI,MO);QuirimbaNationalPark,onInselberg, 8 (holotype:B!;isotypes:BM!,COI!,G!,HBG!, (EA,K,MO);Nampula:24 Oct1965, (K,MO,WAG); António Enes, Dist.Mocambique, (K); António Enes,km5,betweenthelagoon (BR,MO,WAG); Mossuril,Cruce Forest, 18Feb Mozambique: Nóli(dial.macua)( Hexalobus mossambicensis Pedro 5074 MOZAMBIQUE. CaboDelgado:Nexttothe isrepresented byalownumberof Groenendijk 1239 Engl.,Monogr. afrik.Pflanzen- canbedistinguishedfrom the (EA);MocimboabeachDir. velw. Barbosa 2428 H. monopetalus unrbe B ab(iii)(iv). B2 Vulnerable: Pedro 5189 Torre 11944 Groenendijk 1157 (MO,WAG); 0–100m,17 Barbosa 2428 . . (EA);CaboDelgado: (EA,K,MO). ). Thewood is Torre 17348 2 wee these where , , itspresence Gomes eSousa ). ). Gomes Gomes Gomes (COI, an “endangered” one. as recent collections suggesta“vulnerable” statusinsteadof the AOO isjust120 km and oneforest reserve (Ototomo(Cameroon)). Eventhough such asnationalparks(Lopé andMontsDoudou(Gabon)) In addition,severalcollections were madeinprotected areas, herbarium collectionsandhas beenrecently collected (2005). Hexalobussalicifolius May. October andNovember. Fruiting isreported from March to inundated forests andonriverbanks,from sealevelto1,000m. occasionally indeciduouswoodlands,mostlyperiodically secondary tropical lowlandandlowermontanerainforest, (Kouilou region) ( Fig. 10 ). Province), GabonandsouthernpartofRepublictheCongo dark brown, dulltoshiny, rapheraised. Figure 4 I-J. 22–31 mmlong,11–18 mmwide,7–12thick,lightto hairs; seedsca.12–15,biseriate,ellipsoid,slightlyflattened, densely covered withvelutinous, rusty-brown, ca.0.1mm to oblong,6–9.3mmlong,4–5.5cmindiam.,verrucose, the innerandupperside.Monocarps2–4,subsessile,ovoid two lobeswiththeedgescoiledinwards, hairyexcepton tal position,1.2–2mmlong,1.2–1.5wide,dividedin attached tothetopofovaryoftenbendinginhorizon- 3–4, 2–2.5mmlong,1–1.5wide,denselyhairy;stigmas connective appendageflattened,slightlyconvex;carpels narrowly obtriangular, ca.2mmlong,0.5wide, curly, fine,white,ca.1mmlonghairs;stamensnumerous, 0.1 mmlonghairs,baseglabrous orsparselycovered with shorter thanbasalones,insidecovered withappressed, ca. appressed, light-colored, 0.1–0.8mmlonghairs,apicalhairs mm long,4–8wide,rounded atapex,outersidewith ple, tube2–4mmlong,lobesnarrowly lanceolate,17–30 or glabrous; corolla cream toyellow, basalrimdarkpur- covered withcurlyhairs, basalpartofsepalsparselyhairy with rusty-brown, 0.1–0.8mmlong hairs,insidedensely diam.; sepals7–11 mmlong,4–7wide,outsidecovered glabrous; budsovoidto more orlessglobose,ca.7mmin brown hairs,infruit 8–10mmlong, 2.1–3 mmindiam.and 1.8 mmindiam.anddenselycovered withappressed, rusty- above thearticulationcurved,inflower5–8mmlong,1.5– 5–6, ca.5mmlong,3wide;thepartofflowerstalk mm indiam.,oftenbearing1or2vegetativeshoots;bracts articulation 3–7mmlong,ca.1indiam.,fruit 2–2.8 stalk 8–15mmlong;thepartofflowerbelow tinct. Flowers1–3togetheronshortshoots,fragrant;flower at anglesof50–60°withthemidrib;tertiaryvenationindis- veins 5–12pairs,indistinctordistinctabove,belowdistinct, ered withappressed, lightbrown hairsbelow;secondary usually covered withsmall,erect, whitehairsabove, cov- with appressed, lightbrown hairsca.0.2mmlong; midrib glabrous, lowersurfacebrown whendry, sparselycovered tip, basecuneate,coriaceous,uppersurfaceoftengreyish, 5–10 cmlong,1.5–3.5wide,apexacuminatewithablunt brown hairs;leafbladenarrowly elliptictoovateor obovate, long, 1–1.7mmindiam.,sparselycovered withappressed, petioles denselyhairywhenyoung.Leaves:petiole1–3mm quickly turningorange-brown whencut;twigs,leavesand peeling off innarrow, longpaperyflakes,brown togrey, IC Cnevto Status— Conservation IUCN Phenology— Ecology— and Habitat Distribution— Flowering inJanuary, February, May, June, Southern Cameroon (CentralandSouth isrepresented byamoderatenumberof 2 , itspresence inprotected areas aswell A speciesgrowing inprimaryand unrbe B ab(iii)(iv). B2 Vulnerable: Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 12:35:23 2 kmupstream from Akaka camp,10m,14May2005, Maritime: LoangoNationalPark,east sideofRemboNyoungouriver, ca. lage Ekobakona,20Feb1987, chantier SOFORGA,25Mar1989, LeTestu 6387 Ngounié: Mouilaarea, along Boubandjiriver, Mourimatsengui, Feb1927, 1668 31 Jan1966, THE CONGO.Kouilou:road ofBoungolobuildingsite(Pointe-Noire), Bipindi, nodate, Velzen 99 toll postforKribi-Edéaroad, forest atBebamboueII,26May2006, of village,4Jun2006, (P); 21kmNEofKribi,Kribi-Bipindiroad, BidouI,cultivatedfieldsNE 100 m,14Nov2005, m, 14Nov2005, Doudou MountainsNationalPark, ca.5kmSofCampPeny(CBG),100 Yaoundé, 8Jun1933, Nyoma riverbank,12Feb1928, 1663 47 BOTERMANSET AL.: REVISION OFHEXALOBUS 2011] Letouzey8122 ENE ofSangmelima,mapIGN1/200.000Djoum,1,011 m,19Oct1966, Cameroon, 30Oct1991, Kribi, 28 Aug 1970, Wilks 1706 Nov 2005, Mountains National Park,ca.30kmSofMandji,E MontIgoumbi,18 ca. 40kmSofMandji,120m,16Nov 2005, tion thespecimenappeared tobelong Ivory Coast(specimen fang) ( of these twospecieswerefer totheremarks belowthetreatment in agreement with Le Thomas(1969) . Fordifferences between lished. We, however, considerthemastwodistinctspecies, 2000 ), althoughthatcombinationwasneverofficially pub- has beenconsidered asa varietyofthelatter( Wilks etal. Vraua Names— Vernacular Hexalobussalicifolius Uses— Le Thomas(1969) mentionsthepresence at Fig. H. crispiflorus SpecimensExamined— (LBV); Moyen-Ogooué: GoméLake,Jul1952, ); Owoé(dial.yetbou)( 10 Wilks 1706 (WAG); Bipinde,1907, . . (LBV, WAG). Sosef 2376 The fruits (pulp)are edible. (BM,BR,P);Ogooué-Ivindo:laLopéReserve,southof Ayem; Dsrbto o of Distribution Farron 4893 (BR,P);Bidou,nearKribi,28 May1954, Sosef 2266 Zenker s.n. . (WAG); Woleu-Ntem: 33kmEdeMitzic, 13 Apr 1988, ). Sosef 2294 Bos s.n. Unknown s.n. van Velzen 98 (P).GABON.Ogooué,Jun1956, (WAG); 100m,14Nov2005, Hallé 4181 closelyresembles Aké Assi5962 Cameroon: Oouè(dial.jaoudi)( Hexalobus salicifolius CAMEROON. CentralProvince: Vauchi, (F);nodate, Reitsma 3102 (WAG); DoudouMountainsNationalPark, (WAG); Campakok(Camp Akok), South Hédin 1663 Zenker 3330 Médou 1703 McPherson 13813 (WAG); 25kmNofKribi,SEthe (K,P);SouthProvince: 15kmfrom (WAG); HillnearMezese,17km (LBV, MO,NY, WAG); Ogooué- Zenker s.n. ), butaftercloseexamina- (BR);Ototomoreserve, near (BR, G,UPS);Mimfia,near Sosef 2336 ). Gabon:Owui,(dial. . . H. crispiflorus H. crispiflorus (LBV, MO);nearvil- Bois SRF815 (S).REPUBLICOF Harris 8647 H. salicifolius Médou SRFK1703 Sosef 2280 (WAG); Doudou Gauchotte SRFG . (WAG); (LBV); (LBV); Hédin and van in

Cae Plrv, . R B Dumn , n E J Ml 93. 1983. Moll. J. E. in and Drummond , Studies B. R. 2000. K., Palgrave, Maas . Coates M. J. P. and Koek-Noorman, J. W., L. Chatrou, Annonaceae. 1953. M. P. J. Brenan, Caru L W ad . e . 19 tde i Annca XXI. revision A XXXIII. Annonaceae in Studies 1999. He. P. and W. L. Chatrou, 1985. M. H. Burkill, Annonaceaein 35. Fam. 1951. R. in Boutique, 20–29 Pp. Anonaceae. 1862. Hooker . J. D. and G. Bentham, Abnir M 20 2004. M. Arbonnier, Hexalobusbrasiliensis Hexalobusmadagascariensis Aogio, . W J V D Br , n L J G V D Mee 06. 2006. Maesen. D. V. G. J. L. and Burg , D. V. J. W. A., Akoègninou, fieldwork inGabon. permits aswelltechnological,logistic,andbotanicalsupportduring the Issembe, RaoulNiangadouma,andJean-NoëlBoussiengui,forproviding (CENAREST –IPHAMETRA),especiallyLudovicNgokBanak, Yves cle. Finally, wewishtothankthestaff oftheHerbierNationalduGabon for theircommentsandsuggestionsonaprevious versionofthisarti- of distributionmaps.Richard Saundersand Andrew Hippare thanked help related totheuseofBRAHMSsoftware, databasing, andcreation provided nomenclaturalinformation.JanWieringa isalsothankedforhis ture oftheinflorescence. FransBreteler andJanWieringa (WAG) kindly grateful toLubbertWestra, whogavevaluablecommentsonthestruc- who financiallysupportedfieldworkofthefirstauthorinGabon.We are also liketothankthe Alberta MennegaStichtingandMoabiFoundation, of theNetherlands-Leidenbranch)forpreparing thepollenplate.We US, WAG, WRSL,andZ.We are gratefultoBenKieft(NationalHerbarium COI, EA,F, FHO,G,GH,HBG,K,LBV, LISC,M,MO,P, PRE,SRGH,UPS, herbaria fortheloanof,oraccessto,theircollections: A, B,BM,BR,C, Hexalobuscrispiflorus the genus cies. Thesefeatures, among others,pointinthedirection of lianescent habit,charactersnotknownforany covering thefruits andlowersideoftheleafblades, Hexalobus collected inMadagascar. Thesespecimensdonotbelongto type P!),and 347. 1868.Itisonlyknownfrom thetype, Hexalobus give anindicationastowhichgenusitbelongsto. ever, itdoesnotbelongtothegenus observed duringthisstudy. According to Fries (1930) , how- lection oftheMNHN,Paris.Thisspecimenhasnotbeen belongs totheherbariumof A. L.deJussieu,partofthecol- Hexalobusjussiaeanus Species NamesExcludedfrom Hexalobus— Acknowledgments. First describedas The typespecimen,ofwhichthelocalityisnotknown, Africa AnnalsoftheMissouri BotanicalGarden Annonaceae XXXVI.The of Gardens . Botanic Press. Garden Botanical York New York : New 8 . Ruanda-Urundi tarum Montpellier : CIRAD, Margraf publishers,GMBH,MNHN . lytique duBénin sér 2,17:133.1842.= 213, Fig. 5a . 1832.= nudum, herbariumname 338. 1882. 1930) ( Fries, Fusaea . London. . aeon: Sri Publishers. Struik Capetown: becauseoftheirprominent stipes,stellatehairs (?) Uvaria (Baill.)Saff. callicarpus Humbert &Viguier 1044 , d . otqe . Busl . . . . C. A. E. N. I. Brussels : Boutique. R. ed. , . . Lie akus Publishers. Backhuys Leiden: . The usefulplantsofwesttropical Africa Trees, shrubsandlianasofWest-African dryzones Baill., Adansonia 8:348.1868. var. A. St.-Hil.&Tul., Ann. Sci.Nat.Bot., Uvaria We are gratefultothecuratorsoffollowing Brittonia ieaue Cited Literature Baill.,Bull.Mens.Soc.Linn.Paris1: Isolona madagascariensis angustifolia A. DC.,Mém.Soc.Phys.Genève: Duguetia Cardiopetalum calophyllum (?) MemoirsoftheNewYork BotanicalGarden 1 181–203. 51: callicarpa alliance:Where thewayspart. 7 234–245. 87: Hexalobus Rob.&Ghesq.,nomen (P!),bothinfruit and Flore duCongo-Belgeet Baill., Adansonia 8: Chapelier s.n. , buthedidnot Hexalobus (Baill.)Engl. Trees ofsouthern e Royal Kew: Genera plan- Schltdl. Flore ana- (holo- spe- . Copyright (c) American Society for Plant Taxonomists. 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