Copyright (c) American Society for Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Wed, 29 Sep 2021 13:17:12 DOI 10.1600/036364411X553207 © Copyright2011 bythe American SocietyofPlantTaxonomists Systematic Botany ( seven speciesworldwide,withfourintheNewWorld waslastrevised by Noamesi (1958) whorecognized disjunctly distributedintropical Africa and America. The any family().Itconsistsofsmalltolarge trees or Andiroba (Portuguese)belongstothepantropical mahog- Crabwood (English),cedro bateoorcedro macho(Spanish) and Meliaceae oftheNeotropics recognized only and the specimens,thoseof may constituteanecologicalbarriertotheirdispersal.Of all pers. comm.),probably duetodrierhabitatsinthisarea that Amazon, botanical surveyshavebeencarriedoutintheEcuadorian trans-Andean partofthecountry( Fig. 1 examined inthecourseofstudycamefrom thenorthern ). Although many of ( Kenfack 2008 ) suggestedinsteadthatthere are fourspecies acters analyzedinthecontextofamolecularphylogeny vations andamultivariateanalysisofmorphologicalchar- 2007). ( Palacios C.megistocarpa Andes (trans-Andes)inEcuadorandtodayonlytwospecies, tocarpa Gentry (1988) describedthedistinctivecauliflorous treated by Styles (1981) asasynonymof C.nicaraguensis ries with2-ovulateloculi.These specimensare assigned to indumentum onleaves,inflorescences, andflowers, ova- morphospecies comprisesspecimens witharusty farinose branches, were grouped intothree morphospecies. Thefirst The remaining specimens,allwithinflorescences borneon borne onthemaintrunk, andtheirlarge acuminateleaflets. from theothersbasedontheirpoorlyramifiedinflorescences C. guianensis Carapa A recent systematicrevision of Carapa cies ( has cauliflorous inflorescences, provided. late flowersand6-ovulateovaryloculi.Thenewspeciesare described,illustrated,andakey tothefourspeciesrecognized i edges andshortpoorlyramifiedinflorescences, while branches. However, C. nicaraguensis C. surinamensis A. H.Gentry&Dodsonfrom thewesternslopesof Keywords— Abstract— Aubl., commonlyknowninSouth America as C.alticola inEcuador. All Ecuadorianspecimensof Carapa

3 Department ofBiologicalSciences,PontificalCatholicUniversityEcuador, Apartado Postal17-01-2184, (2011), 36(1):pp.124–128 Ab. Aubl., n and , aspeciesdescribed from Nicaraguaand A taxonomicrevision of hasneverbeenfoundthere (DavidNeil, Carapa 4

current address: CenterforTropical Forest Science, Arnold Arboretum, Harvard University, and To e Seis f of Species New Two (butas synonymunder C.DC). Styles (1981) inthetreatment of C. surinamensis C. guianensis, C.alticola cao Meliaceae. Ecuador , , C.longipetala

2 Missouri BotanicalGarden, P.O. Box299,St.Louis,Missouri63166-0299U.S. A. C. megistocarpa 1 Department ofEcologyandEvolutionaryBiologyUniversityMichigan, 2011 KrausNaturalScienceBldg,830N.University Avenue, Ann Arbor, differs from C. nicaraguensis 22 Divinity Avenue, Cambridge,Massachusetts02138,U.S. A. Carapa are recognized inEcuador Miq., ) . Thenewspeciesare closeto Carapa were easilyseparated basedonfieldobser- 5

Author forcorrespondence ([email protected]) C. macrocarpa ai Kenfack David (Meliaceae)inEcuadorisprovided withtherecognition offourspecies, C. guianensis C. guianensis previously describedandcurrently placedassynonymof C. procera omnctn Eio: ea Struwe Lena Editor: Communicating C. guianensis Carapa Michigan 48109-1048U.S. A. C.longipetala C. megis- Ducke, inhavinglarger leafletswithprominent secondaryveins,seedswithrounded Carapa DC). Quito, Ecuador . The . 1 (Meliaceae)FromWestern Ecuador , 2 , 4

, 5 124 C. guianensis n lao . Peréz J. Álvaro and canbedistinguisedfrom different species. morphological discontinuities andthatoccurinsympatry asbelongingto (Kenfack 2011). We considergroups of specimensthatpresent obvious sis ( Gower 1966 ) toexplore the patternsofvariationthesecharacters characters, mainlyfloral,were analyzedusingprincipalcoordinate analy- number ofovulesperlocule.Eighteen quantitativeandfourqualitative inflorescences, andflowers,thelengthofflowerpedicel, andthe the shapeanddimensionsofleaflets, theindumentumofleaves, species suchasthenumberofpairs ofleaflets,thelengthpetiole, acters withemphasisonthoseused inthepasttodistinguishbetween studied inthefield.Specimenswere examinedfor72morphologicalchar- and Esmeraldas. All fourspeciesrecognized herein were collectedand 1990 ). InEcuador, fieldworkwascarriedoutintheprovinces ofPichincha QCNE, STRI,andUS(herbariumabbreviations follow Holmgren etal. specimens from 12herbaria; BR,F, GH,K,MO,NY, P, PANAMA, QCA, Ecuador, Guyana,andPanamaexaminedNewWorld herbarium boundaries in below underthenames Andes. Thesecondandthird morphospeciesare described “Cis-Andean clade”comprisingspeciesthatoccureastofthe west ofthe Andes, while to the“trans-Andeanclade”includingspeciesthatoccur the three Ecuadorianramiflorous morphospeciesbelonged of thenuclearITS(KenfackandColemaninprep.) showthat uli. Furthermore phylogenetic inferences from sequencedata distinctly pedicellateflowersandovarieswith6-ovulateloc- comprises specimensthatdiffer from and ahilumlessthan2cmlong.Thethird morphospecies cences (lessthan20cmlong),andseedswithrounded edges inent secondaryveins,shortandpoorlyramifiedinflores- leaflets withrounded toshortly cuspidateapicesandprom- to subsessileflowersbutdiffer from second morphospeciesincludesspecimensthathavesessile pacific coastfrom Nicaragua toEcuador( Kenfack 2008 ). The C.longipetala geographical rangeof Our findingsare basedonaworldwidereassessment ofspecies basedontheir4-merous flowersborneattheendof Carapa efc respectively. Kenfack aeil n Methods and Material . FortheNeotropics, wecarriedoutfieldworkin 3

C. guianensis C. alticola C. nicaraguensis C. guianensis C. guianensis inhavingdistinctlypedicel- Kenfack& A. J.Peréz and C. guianensis C. guianensis C. megistocarpa s.belongstothe andtwonewspe- extendsalongthe n Ecuadoris inhaving inhaving which Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Wed, 29 Sep 2021 13:17:12 flat. Inflorescence inthe axils offullydevelopedorreduced 8–16 oneachside,networkoftertiaryvenationlooseand rib prominent beneath,glabrous, secondaryveinsprominent, truncate toemarginate, notmucronate, thebasecuneate;mid- lets), glabrous beneath,oblong toobovate,theapexrounded, (basal leaflets)to20–44cmlongand8–13wide(apicalleaf- petiolule 0.8–1.7cmlong;blade13–20long,8–11 cmwide 20–40(–65) cmlong,glabrous; leafletsin4–6(–7)pairs,with in diameter, baseswollen,generallywith2nectaries;rachis 35–60(–100) cmlong,withpetiole12–200.5–1.2 thin, slashpinkish;branchesspreading upwards. Leaves in disturbedforest; buttresses upto2mhigh;barksmooth, and unbranchedhighinprimaryforest, variouslybranched testis laevibus,different. tis veltruncatis usqueademarginata etseminibusrotundatis in foliolisvenissecondariisconspicuisapicibusquerotunda- tiis axillaribusetfloribussubsessilibusvaldeaffine sunt,sed 01 KNAKADPRZ E AAAFO CAO 125 Carapaalticola KENFACK AND PERÉZ:NEWCARAPA FROMECUADOR 1. Indumentumabsentonleaves,inflorescences, andflowers;ovarylocules4-or6-ovulate 1. Indumentumpresent . onleaves,inflorescences, andflowers;ovarylocules2-ovulate 2011] and A mediumsizedtree to30mtall,80cmdbh.Bolestraight Arbores usquead30maltas Fig. . 2. Inflorescence borneonmaintrunk, locules6-ovulate . . 2. Inflorescence inaxilsofexpandedorreduced leaves,locules4-or6-ovulate C. nicaraguensis 78°44 about 5kmfrom Pero Vicente Maldonado,00°00 ECUADOR. Pichincha: Along highwayfrom Quito, R. Valencia. 2150 E. Narvaez,A.Perez, G.Buitron, M.D.Proano, J.Iglesias, 1 3. Flowerpedicel2.5–4mmlong;petals7–9ovarylocules6-ovulate;hilum2–5.5cmlong . 3. Flowerpedicelupto2.5mmlong;petals4–5long,ovarylocules4-ovulate;hilumlessthan2cmlong . . . Dsrbto o of Distribution ′ 40.8˝W, 1,711 m,24Mar. 2007,fl, Kenfack& A. J.Pérez sp.nov.—TYPE: n Ecuador. in Txnmc Treatment Taxonomic (holotype:MO,isotypes:QCA,QCNE). Carapa alticola C. guianensis , , C. longipetala,megistocarpa Key totheEcuadorian Speciesof ininflorescen- D. Kenfack, ′ 52.3˝S, , tane moistforest, 600–2,200 maltitude( Fig. 1 ) September toMarch. 0.5–1.5 mmwide,rounded; testadarkbrown, smooth. Figure 2 . 3.7–6.5 cmwide,upto4pervalve;hilum1.3–2.3mmlong, surface withnumerous nectaries. Seeds3.5–4.7cmlong, 4-valved, eachwithamedianrib,lackingwartyexcrescences, stipe, beakedatapex,green tobrown anddenselylenticellate, cm long,10–15indiameter, globose,narrowed basallyintoa style 1–1.7mmlong;stigmaca1.2indiameter. Fruit 12–20 ical andmore orlesscontinuouswithstyle;locules4-ovulate; yellow. Ovary0.9–1.5mmlongand0.9–1.4diameter, con- long, ca.0.7mmwide.Disk0.8high,2.2–3.2diameter, 8 lobes,lobes1.3–1.7mmlong,straight. Anthers 0.7–1mm with oneortwoglands.Staminaltube4.5–5.5mmlong, Petals 4,4–5mmlong,3–3.5wide,green, glabrous, each brous. Calyx4-lobed,green, glabrous, lobes1.1–1.3mmlong. 4-merous, sessileorsubsessile,pedicel0.8–2.5mmlong,gla- 2 cmlong.Carpellateflowersunknown.Staminate peduncle 2.5–8cmlong,lowermostbrancheslessthan scale-like leaves,lessthan20cmlong,erect, poorlybranched, ers, hasbeenconfusedwith developed orreduced scale-like leavesandsubsessileflow- 79°01 00°08 79°02’W, 00°05’N, 79°02’W, Luteyn 520 del Sr. Lalama,a1,5kmdelsectordeElCristal,00°50’N,78°30’W, Carmen, caminoaChical,00°51’N78°13’W, 79°44’W, of Quinindez,5kmWSantaIsabel,alongoldroad toMono,0°21’N (MO). Esmeraldas:BilsaBiologicalStation,MacheMountains,35kmW Reserva Golondrinas,LasJuntas,00°48’N,78°10’W, Carapaalticola ensis grows onterrafirmeandathigherelevations,while axillaribus etfloribus 4-merissimilesunt,sed infloribuslonge Carapalongipetala from there. in southernColombia,butwehavenotseenanycollections with alonghilumupto3.3cmlong. veins, thefruits are notbeaked,andtheseedsare angular, the leafletsare distinctlyacuminate,withdiffuse secondary with smallerhilumandsmoothseedcoat.In tate fruits, andperhaps mostimportantlyitsrounded and almostunbranchedinflorescences, thebeakedandstipi- larger leafletswithprominent secondaryveins,itsshorter Dsrbto ad Habitat— and Distribution Phenology— Carapaalticola Arbores usquead25maltas AdditionalSpecimensExamined— ′ ′ NY, US,K, QCA,QCNE) D.Kenfack&E.Narvaez2159 01°13 Esmeraldas: Limitofthe ReservaEthnica Awá, prefers inundatedforest andgrows atloweraltitudes. 38.9˝W, 02.2˝N, 79°02 Clark etal.152 Jamarillo 6635 (MO).Pichincha:carretera Quito-PuertoQuito,km113, ′ 46.9˝N, 78°39 Kenfack etal.2155 C differs substantiallyfrom ARAPA Flowering, March to August; fruiting, Brado 429(A) , withitsinflorescences intheaxilsoffully ...... ′ 53.1˝W, (NY);

(MO,QCNE);Parroquia Alto Tambo, frente Finca Kenfack sp.nov.—TYPE: ECUADOR. 3 ...... ′ Kenfack etal.2151 Jaramillo 6970 06.2˝W, 139m,26Mar. 2007, fl, (QCA);Reservaforestal Endesa,00°05’N, (MO,QCA,QCNE). C. guianensis Western Ecuador, inpremon- C. guianense ECUADOR. Carchi: Nortedel (holotype:MO,isotypes: (QCA); ...... Carapa alticola ...... Palacios etal.9689 . However, (QCA);00°06 C. guianensis Jaramillo 7039 ininflorescentiis Vargas etal.4565 C. guianensis C. nicaraguensis C. megistocarpa mayoccur C. longipetala C. alticola C. guian- C. alticola Quelal & byits (QCA); ′ 09.5˝N, (MO);

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Wed, 29 Sep 2021 13:17:12 al. 2150 [Volume 36 SYSTEMATIC BOTANY 126 Fig. 2 ; B, . . Neil 11697

Carapa alticola ; E,F, . A. Leaf.B.Detailofthetertiaryvenation.C.Inflorescences. D.Openedflowers.E.Fruit. F. (basedon: A, C,D, Kenfack etal.2155 ). Kenfack et Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Wed, 29 Sep 2021 13:17:12 D, E,F, 127 KENFACK AND PERÉZ:NEWCARAPA FROMECUADOR 2011] Fig. 3 Kenfack &Narvaez.2159 . .

Carapa longipetala . A. Leaf.B.Detailofthetertiaryvenation. C.Inflorescence. D.Opened flowers.E.Ovary. F. Fruit. G.seed(basedon: A, ; G,H, Kenfack &Narvaez2160 ). B, C, Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Wed, 29 Sep 2021 13:17:12 (MO, QCA,QCNE);nearLita,00°52’N78°29’W, (MO, QCA,QCNE);01°13 Reserva Ethnica Awá, 01°13 indigenous territory, 01°00’N,78°33’W, Cojimíes crossing, 00°19’N,79°55’W, north ofPedernalesalongnewcoastalhighway, 3kmnorthofRío 78°15’W, Awá, Parroquia ElChical,sectorGualpíMedio,ríoCanumbí,01°02’N, ally below600m( Fig. 1 ). December toMarch. (1–)2–3.8 mmwide;testawhitish,smooth. Figure 3 . long, 4–8cmwide,upto6pervalve;hilum2–5.5mm warty excrescences, lenticels andnectaries.Seeds4.3–6.4cm [Volumewidely globose,notbeaked,lightbrown, 4-valved,lackingribs, 36 in diameter. Fruit 18–22cmlong,20–24indiameter, ellipticto flowers, 1.8–2.5mmlonginstaminateflowers;stigma1–1.5 flowers; locules6-ovulate;style0.3–0.7mmlongincarpellate conical andmore orlesscontinuous withstyleinstaminate in carpellateflowers;ca.1mmlongand1.2–1.4diameter, 1.7–2.2 mmlong,2.4–3.3wide,4-locular, 4-ridged,ovoid wide. Disk0.8–1mmhigh,1.2–4diameter, red. Ovary 0.5–0.8 mmwide;antherodes 0.8–1.2 mmlong,ca.0.6 1.7–2.2 mmlong,notreflexed. Anthers 1.2–1.5mmlong, glabrous. Staminaltube6–7.5mmlong,with8lobes,lobes 1.3–1.8 mmlong.Petals4,7–9long,4–7wide,green, 2.5–4 mmlong,glabrous. Calyx4-lobed,green, glabrous, lobes 2 cmlong.Flowers4-merous, distinctlypedicellate,thepedicel SYSTEMATIClong, erect, peduncle2–11 BOTANY cmlong,lowermostbranchesupto axils offullydevelopedorreduced scale-likeleaves,18–45cm tiary venationlooseandinconspicuous.Inflorescences inthe glabrous, secondaryveins9–23oneachside,networkofter- not mucronate, thebaserounded; midribprominent beneath, pairs, glabrous beneath,oblong,theapexshortlycuspidate, leaflet pairs;25–72cmlongand9–20wideinultimate 1–1.5 cmlong;blade15–27long,7–13wideinbasal 30–48 cmlong,glabrous; leafletsin5–7pairs,withpetiolule in diameter, baseswollen,generallywith2nectaries;rachis Leaves 50–75cmlong,withpetiole15–300.7–0.9 buttresses lowto1mtall;barkwhitish;branchesspreading. gle stemmedwhenyoung,becomingbranchedwithage, ulis 6-ovulatis,different. pedicellatis, foliolisapicibusbreviter cuspidatisetovariisloc- 128 Distribution— Phenology— A smalltomedium-sizedtree to25mtall,30cmdbh,sin- AdditionalSpecimensExamined— Grijalva etal.503 Flowering, March toSeptember;fruiting, Western Ecuador, inlowlandforest, gener- (MO); ′ 46.9˝N, 78°39 ′ 54.8˝N, 78°39 Rubio etal.1681 ECUADOR. Carchi: ReservaEthnica Neill &QCNE11697 Ortiz etal.123 ′ 06.2˝W, ′ 26.9˝W, (MO).Esmeraldas:50km Kenfack &Narvaez2159 Kenfack &Narvaez2158 van derWerff etal. (NY);limitofthe (O; Awá (MO);

sis six pervalve,are more angularwhilethoseof smooth seedcoats.Howevertheseedsof raguensis et al.173 Naei G K. 15 1958. K. G. Noamesi, 2008. D. Kenfack, 9501 Sye, 91. Sitnoda P 5 2 n in 359–420 Pp Swietenioideae . 1981. B. Styles, 00°52’N 78°26’W, Plco, . 07. 2007. A. W. Palacios, Herbariorum . Index 1990. Barnett . C. L. and Holmgren , H. N. P., meth- Holmgren, vector and root latent of properties distance Some 1966. C. J. from Gower, for combination new a and species New 1988. H. A. Gentry, trations are by A. Arbelaez. and laboratoryworkbytheCenterforTropical Forest Science.Theillus- with logistics.FieldworkwasfundedbytheNationalGeographicSociety D. Neil,M. Asanza andE.Narvaezfortheirhospitalityhelping J. Iglesiasfortheirhelpwiththeexportationpermitandduringfieldwork; Dr. RenatoValencia andhis“Carapateam”,G.Buitron, M.D.Proano, and herbaria BR,F, GH,K,NY, P, andUSforproviding materialonloan; sympatric with refers to (1988) inwesternEcuadorashavingorangenectariesclearly The populationof disk) unlike Tirado etal.287 Narvaez 2160 Knak, D . 21 eurcin n in Resurrection 2011 . D. Kenfack, C.guianensis and have sessiletosubsessileflowers.Furthermore, tinctly pedicellateflowerswhile Acknowledgments. Carapalongipetala , onlytwopervalve,haverounded edges. (MO);reserva ethnica Awá, 01°14 Tyo e Yr h Nw ok oaia Garden . Botanical York New The York : New Taylor . No 28: Swedish Research CouncilPubl.House. Mdsn: Uiest o Wisconsin . of University Madison: http://www.nybg.org/bsci/ih/ . Garden , Botanical York New ed. 8th ods usedinmultivariateanalysis . 1429–1439. 75: trans-Andean South America . Society methods inaphylogeneticcontext. morphological variationandspeciesboundariesusingmultivariate sertation. St . Louis : University ofMissouri-St.Louis. C. alticola (MO,QCNE); are comparableinsizeandbothhavewhitish C. longipetala 15 186–221. 165: Meliaceae (MO,QCA,QCNE);centro Guadualito,01°43’N,78°53’W, C. longipetala, inwesternEcuador. (MO);01°15’N78°40’W, havewhitishfloralnectaries(alsoknownasa Systematics andevolutionof Aulestia &Aulestria1427 C. longipetala A revision ofXylocarpae(Meliaceae) , d. T D Pnigo , .. tls ad . . H. A. D. and Styles, B.T. Pennington, D. T. eds. , 98. , like Aulestia etal.227 The authorsare gratefultothecuratorsof . Theseedsof Meliaceae ieaue Cited Literature C. alticola which hasorangetored nectaries. intheReservaEthnica Awá. Carapa Annals oftheMissouriBotanicalGarden . FloraofEcuador Biometrika C. guianensis ′ 46.9˝N, 78°42 Aulestia etal.105 (Meliaceae):areassessment of Carapa longipetala (MO,QCNE). Botanical JournaloftheLinnean , hasbeenconfusedwith (MO);Parroquia Alto Tambo, C. longipetala Carapa mentionedby Gentry Flora Neotropica Monograph 3 325–33b8. 53: C. longipetala (Meliaceae). C. megistocarpa . Ph.D.dissertation o 2. Stockholm: 82. No ′ 42.6˝W, and C. nicaraguen- (O; (MO); C. guianensis n and C. alticola hasdis- Ph. D.dis- Kenfack & C. nica- u to up , Aulestia is