Systematic Wood Anatomy of the Pavetteae (Rubiaceae, Ixoroideae) Author(s): Steven Jansen, Petra De Block, Hans Beeckman and Erik Smets Source: Systematics and Geography of Plants, Vol. 68, No. 1/2, Morphology, Anatomy and Systematics at the Centenary of Wilhelm Troll's Birth (1999), pp. 113-133 Published by: National Botanic Garden of Belgium Stable URL: http://www.jstor.org/stable/3668595 . Accessed: 26/03/2013 05:06 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. . National Botanic Garden of Belgium is collaborating with JSTOR to digitize, preserve and extend access to Systematics and Geography of Plants. http://www.jstor.org This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr.P1.68: 113-133(1999) woodanatomyofthePavetteae Systematic (Rubiaceae,Ixoroideae) StevenJansena*, PetraDe Blockb,HansBeeckmanc & ErikSmetsa ofPlantSystematics, BotanicalInstitute, Kardinaal Mercierlaan K.U.Leuven, 92,B-3001Heverlee, Laboratory Belgium forcorrespondence *author [Steven.Jansen @bio.kuleuven.ac.be] ofBiology, ofAntwerp & National BotanicGardenofBelgium, University b Department DomeinvanBouchout, B-1860Meise,Belgium c Africa, 13,B-3080Tervuren, RoyalMuseumforCentral Leuvensesteenweg Belgium - Woodanatomy Abstract. of16 generaofthePavetteaehas beenexamined. Exceptforsomegeneral such as and thefrequent the Ixoroideae, solitaryvessels,presenceoffibre-tracheids, featuresof was cell variation occurrenceofdisjunctive rayparenchyma walls,diagnostic foundinaxialparenthe attention was to associaand morphology ofvestures features.Special given chyma, rays, crystal an Ixora ted withvesselpitsand twovesturetypesweredefined.Thedelimitation of groupis wood or lines axial the small bands of parenchyma, styloidsor anatomicallysupportedby presenceof and vestures that uniseriate branched rays, completely fillthepit predominantly elongatecrystals, is in theray are derived. Wood that these characters chamber.It is suggested ofDuperrea different this an corroborates exclusion andthepresenceofremarkable bands; from parenchyma composition inchambered axialparenchyma cellsoccurmainlyin thePavetteae. crystals Longchainsofprismatic butare also presentinNichalleaand Pavetta. Bridson'sinformal groupI ofTarenna, woodanatomy. Keywords.- Rubiaceae,Ixoroideae,Pavetteae,Ixoreae,systematic Introduction ? 20 generaand? 1200species Rubiaceaeandcomprise The Pavetteaearea largetribeofthefamily andSouthAmerica.Being Ixora reaches Central are table (see 1). They essentially only paleotropical; thetribe are well of interest, although species Ixora undoubtedly known, plantsofhorticultural tropical ornamentals. other containsseveral potential offlowers incertain areveryvariable ThePavetteae characters, (1e.g.thenumber perinflorescence oftheleafblade(1-50cm).Furthermore, ofthecorollatube(0.3-20cm),andthelength 400),thelength a character ofovulesperloculevariesfromonetonumerous, thenumber heavilyrelieduponbyearly theRubiaceae,grouping threeinformal serieswithin Hooker(1873) recognized taxonomists. species was takenoverbySchumann ormanyovulesperlocule.His classicalsystem withone,twocollateral theCinchonoideae Thelatter, however, (1891) withfewmodifications. onlytwosubfamilies, recognized and Coffeoideae with one ovule ovule locule the than one more characterized perlocule.As a per by two subfamilies inaccordscattered over the Pavetteae were now constitute the the which result, genera Allrights reserved. Subjecttocopyright. Permissionforuse mustalways be obtainedfromtheNationalBotanicGardenof Belgium. BotanicGardenofBelgium ? National This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions ISSN 0303-9153 Syst.Geogr. P1. 68 (1999) Table1.Generastudied. 1985. followsRobbrecht Tribalcircumscription 1984,1988andBridson& Robbrecht species genus distribution wood samples Captaincookia N.Halle CladocerasBremek. 1 NewCaledonia 1 1 EastAfrica (Kenya,Tanzania) 1 ColeactinaN.Hall6 1 Gabon& Congo Hook.f. Dictyandra 2 Africa West& Central tropical 3 DoriceraVerdc. 1 (Mascarenes) Rodrigues 2 DuperreaPierreex Pitard ?2 Indo-China 1 HomolleaAr6nes ?3 Madagascar 1 Homolliella Ar~nes ?5 Madagascar IxoraL. + 400 pantropical 12 Hook.f. Leptactina + 25 Africa tropical 10 Comm.ex Juss. Myonima 4 Mascarenes 1 NichalleaBridson 1 Africa West& Central tropical 5 K.Schum. Pachystylus 1 NewGuinea 1 Madagascar PavettaL. 5 + + 400 paleotropical 20 RutideaDC. + 20 Africa tropical 8 ParacephaelisBaill. Homolle Schizenterospermum ex Arenes 4 Madagascar paleotropical 14 1 Kenya& Somalia 1 4 NewGuinea,SolomonIslands 5 TarennaGaertn. + 180 TennantiaVerdc. Val. Versteegia ofthegenerastudiedsee table2). Dictyandra, ancewiththeirovulenumber (forthetaxonomic history oftheCinchonoideae tobe members (tribeGardenieae), LeptactinaandTarennae.g.wereconsidered whileuniovulate (tribeIxoreae).More generasuchas IxoraandPavettawereplacedintheCoffeoideae suchas Bremekamp (1988) (1958), and Robbrecht (1952, 1966), Verdcourt recentlytaxonomists whichresultedin morenaturalsystems. consideredthenumberof ovulesperloculeless important, threesubfamilies Verdcourt (1958) recognized mainlybasedon thepresenceor absenceofraphides, theCinchonoideae and seeds albuminous), (raphidesabsent, namelytheRubioideae(raphidespresent He placedthe seeds and theGuettardoideae seeds albuminous), + exalbuminous). (raphidesabsent, these recent more in same and Ixoreae the classifications, twotribes Gardenieae subfamily. Following Robbrecht to the Ixoroideae 1988, 1966, 1994). subfamily (Bremekamp belong Alreadyin 1934 BremekampsuggestedthatwithintheIxoreae the groupwiththe terminal is a natural one.Robbrecht inflorescences (1984) revivedforthisgroupthenamePavetteaeA.Rich.ex the inhistribethosegeneraoftheold Gardenieae-Coffeeae andincluded Dumortier complexcombining seeds with one to small terminal characters: inflorescences, many freely relatively drupes following 114 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions P. De Block,H. Beeckman& E. Smets,Woodanatomy ofPavetteae S. Jansen, (Rubiaceae) absentor exotestalcells withthickenings theplacenta,seedswithan adaxialexcavation, surrounding tectate and the outer wall, 3-4-colporate pollengrains. tangential occurring along ofthePavetteaehasnotbeenquestioned. Untilrecently, thedelimitation However,De Block& Robthe brecht(1997a) excludedtheIndochinesegenusDuperreawhileAndreasen(1997) transferred tribeIxoreae. generaIxora,Doricera, Captaincookia,Myonimaand Versteegiato thenewlyresurrected Table 2. TaxonomichistoryofthePavetteae. Author Genera Tribe(subfamily) Schumann1891 Ixora,Myonima, Pavetta,Rutidea Ixoreae(Coffeoideae) Pachystylus, Tarenna Gardenieae Dictyandra, Leptactina, Paracephaelis, (Cinchonoideae) Ixoreae(Ixoroideae) Homollea, Homolliella, Ixora, Duperrea, Bremekamp Pavetta, Paracephaelis, Pachystylus, 1934,1952,1966 Myonima, Tarenna Rutidea,Schizenterospermum, Coleactina, Dictyandra, Leptactina Cladoceras,Versteegia Gardenieae (Ixoroideae) uncertain position Pavetteae(Ixoroideae) Cladoceras,Coleactina, Captaincookia, Homollea, Doricera, Duperrea, Dictyandra, Homolliella, Ixora,Leptactina, Myonima, Pavetta, Nichallea, Paracephaelis, Pachystylus, Tarenna, Rutidea,Schizenterospermum, Tennantia, Versteegia Andreasen1997 Cladoceras,Coleactina, Homollea, Pavetteaesensu Dictyandra, Nichallea, Homolliella, Pachystylus, stricto(Ixoroideae) Leptactina, Pavetta,Rutidea, Paracephaelis, Tarenna,Tennantia Schizenterospermum, Robbrecht1984, Bridson& Robbrecht1985, Robbrecht1988 Doricera, Ixora,Myonima, Captaincookia, Versteegia Ixoreae(ixoroideae) Duperrea Gardenieae (Ixoroideae) oftheRubiaofrepresentatives Thepresentstudyis partofa seriesofwoodanatomical investigations ceae. In additionto theworkof Koek-Noorman 1969a, 1969b, 1972, 1977), our (Koek-Noorman aim at a globalsurveyof thewood anatomicalvariationof thislarge, observations complementary ofthe thedelimitation dataofthePavetteaeareusedtocorroborate Wood anatomical tropicalfamily. atthe13th A posterofthispaperwas presented infratribal tribePavetteaeandtoclarify relationships. etal. 1997a). andSystematics (Jansen Anatomy Symposium Morphology, Material,methodsand terminology herbarium thexylarium ofTervuren Allwoodsamplesaretakenfrom (BR,K, L, P) orfrom (Tw),from livingcollections specimens totheirorigin, examined arelistedbelowwithreference Thespecimens inthegreenhouses oftheNational Botanic GardenofBelgium. ofthewoodsampleifknown. collector anddiameter followed forSEM. Themethods forLM and/or cleancutsurfaces toobtainsections Thewoodwascutwitha slidingmicrotome whichareindicated etal. (inpress).Woodof29 specimens aredescribed below,was investigated bySEM. byan asterisk byJansen The SEM sampleswerecoatedwithgoldand The micromorphology of vestured pitswas onlyobservedin these29 specimens. at27 kV. electron examined witha JeolJSM-6400scanning microscope forhardwood identification features followstheIAWAlistofmicroscopic 1989).Cellswhich Terminology (IAWACommittee to vesselmembers' so called'fibriform resemble fibres buthaveoneortwoverysmallperforations, 1935),arereferred (Woodworth becausetheyareperforate. as narrow vesselelements 115 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr. Pl. 68 (1999) Listofspecimensstudied Captaincookia margaretaeN.Hall6: New Caledonia: MacKee24542(P!), 11mm* Pouembout, Cladocerassubcapitatum (K.Schum.& K.Krause)Bremek.: Harris4464(K!), 5 mm* Tanzania: Pandeforest reserve, arborescens Welw.ex Benth.& Hook.f.:Congo(Dictyandra Tw 2416,> 10cm*; voucher Maudoux Kinshasa):Kisangani, Michelson1332 1887 (BR). Rwanda: Tw 23511; voucher (BR). Angola:CuanzaNorte,Tw 28571,> 10cm*; voucher Murta& Da Silva 1526(BR) Dechamps, Doricera trilocularis (Balf.f.)Verdc.:Rodrigues:Grande Coode4316 (K!), 5 mm; Solitude,Tirvengadum Montagne, 908 (K!), 5 mm* Duperrea pavettifolia(Kurz) Pitard:National Botanic Garden of Belgium: cultivationnumber 1000-7777 8 mm* exactoriginunknown), (Indochina, Homollea longiflora Arhnes: Madagascar: Majunga, Tw 44766;voucher Dorr& Koenders 3017(P!) Ampijoroa, Ixora brachypoda DC.: Congo(-Kinshasa): Yafunga, LusambilaRiver,Tw 33739; voucherLouis 4147 (BR!); Louis7197(BR!); nearIsalowe,Tw 38466;voucher Yafunga, Tw 33700;voucher Louis Yamboa,25 kmNWofYangambi, 3626(BR!) Ixora ferrea Benth.:Guadeloupe: Tw 47667, 25 mm*; voucher Rollet1147(n.v.) Tw Ixora foliosa Hiern:Cameroon:CameroonMountain, MC 170(K!) De Wilde& Letouzey Breteler, 13222;voucher Ixora gracilifloraBenth.:Surinam: Tw 35188; voucher Lindeman5012 (n.v.) Ixora laxifloraSm. var.laxiflora:Liberia: GrandBassa Tw 26695;voucher DukwiaRiver,Monrovia, Cooper County, 307 (FHO!) andTw 30340;voucher Cooper308 (FHO!) Ixora mildbraedii K.Krause:Congo(-Kinshasa):Yangambi, Tw 33413;voucher Louis 1381(BR!) Ixora nigricans Zoll. ex Miq.:India: Tw 44916,> 10cm*; voucherMaharashtra s.n. (n.v.) Ixoraperuviana(Spruceex K.Schum.)Standl.:Brasil:Acre, RioPurus,Tw 34425;voucher (n.v.) Krukoff5590 Ixora sp.: Peru: San Martin, Tw 29340,35 mm*; voucher Williams6717 (n.v.) LeptactinaarnoldianaDe Wild.:Congo(-Kinshasa):Tw Donis2163(BR) 32476,18mm*; voucher Leptactinaleopoldi-secundi Congo(-Kinshasa):Tw Btittn.: Louis6403(BR) 38285,10mm*; voucher manniiHook.f.:Congo(-Kinshasa):Kouilou,Tw Leptactina 49066,10mm;voucher Dechamps13065(BR) LeptactinapynaertiiDe Wild.: Congo(-Kinshasa):Tw Louis3983(BR) 35582;voucher Hook.f.:Guinea:Lab6,Tw 47182, Leptactinasenegambica 17 mm *; voucherStassarts.n. (n.v.) LeptactinaseretiiDe Wild.:Congo(-Kinshasa):Tw 33276, 10 mm*; voucherLouis 1794 (BR); Tw 34977; voucherLouis 3194 (BR); Yangambi,Louis 1507 (BR), 13 mm * Leptactina surongaensis De Wild.: Congo(-Kinshasa): Tw 35830; voucherLouis 6045 (BR) Myonimaviolacea(Hiern)Verdc.:Mauritius:Friedmann 1198(P!), 6 mm* Nichallea soyauxii(Hiern)Bridson:Congo(-Kinshasa): Louis Yangambi,N'Gaziroad,Tw 32837,22 mm*; voucher Tw 33267;voucher 429 (BR!); Yangambi, Louis1778(BR!) ; Yangambi,N'Gazi road,Tw 34928; voucherLouis 2963 N'Gazi road,Tw 34958; voucherLouis (BR!); Yangambi, 3030(BR!) K.Schum.: guelcherianus Pachystylus Papua-NewGuinea: Sands1074(K!), 11mm* Amanab, Kilifas, PavettabilineataBremek.:Congo(-Kinshasa):Shaba,Tw Malaisse11318(BR) 40961;voucher Pavettabrachycalyx (Hiern)Bremek.:Congo(-Kinshasa): Tw 32995; voucher Louis832 (BR); Tw 33222,20 mm*; Tw 38298; voucher voucherLouis 1540(BR); Yangambi, Louis6453(BR!) Pavettacalothyrsa Bremek.: Congo(-Kinshasa):Yangambi, Tw 33079;voucher Louis 1017(BR!); Yangambi, Isalowe, Tw 35799;voucher Louis5944(BR!) cellulosaBremek.: Pavetta Congo(-Kinshasa):Bantoie,near Louis2181(BR!) Eala,Tw 33397,*; voucher PavettacrassipesK.Schum.:Burundi: Bubanza,fermede Lewalle1361(BR!); Randa,Tw 17248,10 mm*; voucher Tw 32020; voucher Congo(-Kinshasa):Shaba,Fungurume, Malaisse9450(BR!); Katanga, Tw 40953;voucher Dikulushi, Malaisse11331(BR!) Pavettagardeniifolia Hochst.ex A.Rich.var.gardeniifolia: Tw 40966; voucher Luiswishi, Congo(-Kinshasa):Katanga, Malaisse11541(BR!) Pavetta nitidulaWelw.ex Hiern:Angola: Malanje,Tw Murta& Da Silva 1488 28361,18mm*; voucher Dechamps, (BR!) Pavettaoblongifolia (Hiern)Bremek.: Senegal:Casamance, De Wolf99(n.v.) Tw 52589;voucher Pavetta puberula Hiern:Congo(-Kinshasa): Luki, Tw Donis2086(BR!); Luki,Tw 32587;voucher 32428;voucher Donis2335(BR!) Pavetta schumannianaHoffm.F.ex K.Schum.:Angola: Cunene,Tw 28658; voucher Dechamps,Murta& Da Silva 1298(BR!);Congo(-Kinshasa): Tw 23613andTw Luiswishi, Malaisse6712 (BR!); Congo(-Kinshasa): 23614; voucher Tw 29052;voucher de Witte 4687(BR) Haut-Lomani, Rutidea fuscescens Hiern subsp.fuscescens: Congo(Tw 40964; voucher Malaisse Kinshasa): Shaba,Luiswishi, 11542(BR!) RutideahispidaHiern:Congo(-Brazzaville):Kouilou,Les Sara,Tw 49272;voucher Dechamps13211(BR!) Rutidea Hiern:Congo(-Kinshasa):Tw 31769, membranacea 8 mm*; voucher Girard5188(BR) RutideasmithiiHiern:Congo(-Kinshasa):Bokali,Befale, Tw 40352,30 mmn *; voucher Dechamps8052 (BR!); Luki, Tw 32436; voucherDonis 2097 (BR!); ibid.,Tw 32591: voucherDonis 2341 (BR!); Yangambi,N'Gazi road,Tw km 13 on 34914; voucherLouis 2947 (BR!); Yangambi, Louis7467(BR!) N'Gaziroad,Tw 38504;voucher Tarennaattenuata Tw (Hook.f.)Hutch.:China: Guangdong, 42126, > 10 cm *; voucherForest Inst. 1546 (n.v.) 116 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions S. Jansen, P. De Block,H. Beeckman & E. Smets,Woodanatomy ofPavetteae (Rubiaceae) Tarenna bipindensis K.Schum.: Congo(-Kinshasa): Plateau,Tw 38302, 13 mm*; voucher Yangambi,Itasukulu Louis6462 (BR!) TarennaeketensisWemh.:Congo(-Kinshasa):Tw 35792; voucher Louis5918(BR) K.Schum.: Tarennafusco-flava Congo(-Kinshasa):Luki,Tw Donis2431(BR!); ibid.,Tw 41352,10mm*; 32654;voucher voucher 2387(BR!) Wagemans Tarenna graveolens(S.Moore) Bremek.:Tanzania: Tw 39207 Tarenna neurophylla (S.Moore) Bremek.: Congo(Kinshasa): Katanga,Tw 45140; voucherMalaisse 13411 (BR!) Tarennapavettoides Bridson: (Harv.)Sim.subsp.gillmannii Tw 44603; voucher Congo(-Kinshasa):Shaba,Luiswishi, Malaisse13425(BR!) Tw TarennapetitiiN.Hall6:Congo(-Kinshasa):Yangambi, Louis5992(BR!); Yangambi, Luweo 35821,10mm;voucher Louis7134(BR!) Plateau,Tw 38452;voucher Tarenna precidantennaN.Hall6: Congo(-Kinshasa): Yangambi,LuweoPlateau,Tw 38479; voucherLouis 7258 (BR!) Tarenna supra-axillaris (Hemsl.)Bremek.:Madagascar: Tw 44778, 17 mm*; voucher Dorr& Majunga,Ampijoroa, Koenders2966(P!) Tennantia sennii(Chiov.)Verdc.& Bridson: Kenya: Isiolo, Kibui& Mungai5661(K!), 7 mm* Gilbert, caulifloraVal.: New Guinea: Baiteta,Madang, Versteegia RidsdaleNGF33904(L!), 7 mm minorVal.: New Guinea: Beaufort River,Pulle Versteegia 348,(L!), 5 mm solomonensis Ridsd.:SolomonIslands:N.E. San Versteegia Whitmore 6161 (L!), 19 mm;Northof Tetepari Christobal, MauriasiBSIP15875(L!), 8 mm* island, Versteegia sp.: New Guinea: Okapa,Awande,Katik62190 (K!), 6 mm Results inDictyandra todistinctly absentbutindistinctly Growthringsaregenerally arborescens(Tw present seretii Louis Tw 2416, (Tw 33276, 28571), Leptactina 1507), Myonimaviolacea,Pachystylus Pavettabilineata,Rutideafuscescens,R. membranacea, Tarennaneurophylla, T. guelcherianus, and solomonensis BSIP (Mauriasi 15875).Theyareformed (Tw 44603), Versteegia pavettoides byan a decreaseintheamount ofvessels,a changeinvesseldiameter increaseinthefibrewallthickness, or thepresenceofmarginal parenchyma. of 2-3(4) vesselsoccurin and solitary Vessels are diffuse (80%-90%) (fig.1-9). Radialmultiples in Tw 40352 ofRutidea of arborescens 40 Tw 23511 %, fig.10), specimen (ca. Dictyandra specimen 80 and in Tarenna smithii Vessel outlineis oval, 50 %, fig.12). %, fig.11), (ca. (ca. supra-axillaris Tw Vessel Pavetta are circularor somewhat brachycalyx 33222,fig.3). perforations irregular (e.g. in that show vestures are found arnoldiana Leptactina frequently alwayssimple.Simpleperforations (ca. 80%,fig.37) andPachystylus guelcherianus. < 50 pm.Verysmallvessellumina diameter ofvesselluminais rather Thetangential small,mostly > 60 and occurin Captaincookiamargaretae Duperreapavettifolia (fig.9). Vesselswitha diameter seretii(Tw 33276,Louis 1507,fig. inDictyandra arborescens (Tw 2416),Leptactina pmarepresent (Tw 41352,fig.4). The 1), Rutideasmithii (Tw 32591,Tw 40352,fig.11),andTarennafusco-flava Pavettabrachycalyx ofvesselspermm2is on average70 but> 100inDuperreapavettifolia, number is relatively low in all speciesof (Tw 33222), P. cellulosa,and Tennantiasennii.Vesselfrequency L. seretii(Tw 34977),Rutideahispida,R. membranacea, Ixora(fig.5,6),Leptactina leopoldi-secundi, classes(about30 and 80 pm) occurin and R. smithii (Tw 40352, fig.11). Vesselsoftwodiameter and in all speciesofRutidea.Vessel elementlengthis Tarennafusco-flava(fig.4), T. eketensis, arborescens(Tw between 250 pmand 1250pm,on average700 pmn. TylosesarefoundinDictyandra 2416). areoftenpresent inspeciessuch withoneortwoperforations Notethatverysmallvesselelements seretii(Louis 1507,Tw Ixorabrachypoda as Duperreapavettifolia, (Tw 33700,fig.53), Leptactina (Tw 32591) andPavettacellulosa. 34977),Rutideasmithii 117 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions nill lp t..0 * fTnrW I,.,-4p W m5..6 ...-J~ .. rr ...'- -._ " .+'I - - _ Ilia,! [] ml . , _74 j. _- b m,m--_?- ofaxial parenchyma:1, Leptactinaseretii(Tw 33276), Figure 1-9. Transversesectionsshowingsolitaryvessels and distribution axial parenchymascarcelypresent(bar = 100 pm); 2, Leptactinaarnoldiana,axial parenchymadiffuse(bar as infig. 1); 3, Pavettabrachycalyx(Tw 33222), axial parenchymadiffuse(bar as infig. 1); 4, Tarennafusco-flava(Tw 41352), relativelywide vessels and axial parenchymadiffuse(bar = 100 pm); 5, Ixora nigricans,axial parenchymadiffuse(bar as infig. 4); 6, Ixora withtwoparenchymabands (bar as infig. 4); 7, Doricera trilocularis(Coode ferrea,axial parenchymadiffuse-in-aggregates 4316), axial parenchymabands 3 cells wide (bar = 100 pm); 8, Versteegiaminor,parenchymabands present(bar as infig. 7); 9, axial parenchymadiffusewithunusualparenchymabands (bar = 100 pm). Duperreapavettifolia, This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions _ -A maim 4 4 -0 , Figure 10-18. Transverseand tangentialsections: 10, Dictyandraarborescens(Tw 23511), vesselsfrequentlyin radial multiples and axial parenchymadiffuse(bar = 100 pm); 11, Rutideasmithii(Tw 40352), wide vessels in radial multiplesand wide,very vessels in radial multiplesand axial parenchymain thin-walledfibre-tracheids(bar as infig. 10); 12, Tarennasupra-axillaris, aggregates (bar as infig. 10); 13, Dictyandraarborescens(Tw 2416), 1-4-seriaterays (bar as infig. 10); 14, Ixora ferrea:rays uniseriateor biseriateover a shortlength(bar as infig. 10); 15. Versteegiasolomonensis,raysuniseriate,rarelybiseriate(bar as multiseriate infig. 10); 16, Duperreapavettifolia,multiseriaterays mainlyupright(bar = 100 pm); 17, Tarennaneurophylla, fibres(bar as infig. 16); 18, Leptactinaseretii(Tw 34977), uniseriaterays (bar as infig. 16). raysand septatelibriform This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr. Pl. 68 (1999) roundtooval,andminute Vessel pitsarealternate, (2-4 gsm).PitsinMyonimaviolacea(fig.32) are inLeptactina occur seretii(Tw 34977,Louis 1507), Non-bordered often moreelongated. vessel-ray pits butnon-vestured andTarennasupra-axillaris.All vesselpitsarevestured, pitsareobservedinPavetta Two typesofvesturescouldbe distinguished in the cellulosa(fig.19) andTarennasupra-axilaris. SEM. examined by species fillthepit vestures occuraroundthepitcanalandonlypartly Type 1: smallandusuallyunbranched Vesturedpitsof thistypeare foundin chamber.This typeneveroccludestheouterpit aperture. arborescens(Tw 2416,Tw 28571,fig.21), Ixoranigricans, Cladocerassubcapitatum, Dictyandra L L seretii(Tw 33276,Louis 1507),Nichalleasoyauxii(Tw arnoldiana, leopoldi-secundi, Leptactina (Tw 33222), P. crassipes(Tw 17248),P. nitidula(fig.22), Rutidea 32837), Pavettabrachycalyx membranacea,Tarennaattenuata(fig.24), T. bipindensis (Tw 41352), and (fig.23), T.fusco-flava Tennantiasennii(fig.20). fillthepitchamberandoccludetheouterpitaperture. The vestures completely Type 2: numerous intoa rather arebranched andoftenfasciated in Captainvestures compactnetwork. Type2 is present cookiamargaretae 908,fig.27),Duperreapavettifolia (Tirvengadum (fig.26), Doriceratrilocularis violacea,Pachystylus guelcherianus (fig.28), Versteegia cauliflora (fig.29), Ixoraferrea,Myonima violacea Myonima Specieswiththisvesture type,suchas Ixoraferrea, (fig.25),and V.solomonensis. and V.solomonensis (Mauriasi cauliflora, (fig.32), Pachystylus guelcherianus (fig.31), Versteegia showvestures ontheinnerpitaperture. BSIP 15875),also frequently thatforsomespecies,e.g.Ixorasp. andTarennaattenuata(fig.24), the We shouldremark, however, inthe Thelatter betweenbothtypesis somewhat distinction arbitrary. speciespossessesmorevestures in T. attenuata toother specieswithvesture type1 (fig.20-23),butthevestures pitchambers compared whichis characteristic fortype2. Ixorasp. appearsto do notcompletely occludetheouterpitaperture be intermediate betweenbothvesturetypesas notall outerpitapertures areoccluded.We classified arebranched andcompletely vestures ofDuperreapavettifolia (fig.29) intype2 becausethevestures occludetheouterpitaperture. However,theyappearto be moreregularandbead-likethannormal in LeptactinasenegambicaandRutidea oftype2 (fig.25-28). The vestures present representatives fromthetypesdescribed smithii above:theyarebranched andanastomosed, in (Tw40352)aredifferent theouteraperture, buttheydo notprojectintothepitchamber. partoccluding In species withvestured innerpitapertures vestured (fig.30-32),thevesselwallsaresometimes aroundthepitapertures as observedin Ixoraferrea,Pachystylus guelcherianus (fig.31), Rutidea smithii Note that we no in and found differences vesture (Tw40352), Versteegia cauliflora. morphology betweendifferent typesofvesselpits(intervessel pits,vessel-ray pits,vessel-axial parenchyma pits,or vessel-fibre pits). Fibresarenon-septate. tothetangential bordered andradialwalls.Outerand Distinctly pitsareconfined often showvestures whichareverysmallandsometimes inner pitapertures scarcely (fig.33-36). present Within thesamespecimen arborescens Tw 28571),fibrepitscanbe slightly vestured (e.g.Dictyandra or non-vestured. arefoundon theinnerfibrewallsofRutideasmithii Distinctvestures (Tw 40352). bordered inLeptactina seretii(Tw 34977) and Septatefibreswithsimpletominutely pitsarepresent Tarennaneurophylla withsimplepitsconfined fibres totheradialwallsoccurin (fig.17). Non-septate Tarennasupra-axillaris. Thefibres arethintothick-walled. seretii(Tw 34977) VerythinfibrewallsarefoundinLeptactina andRutideasmithii (Tw 40352,fig.11).Thefibrewallsinthelatter speciesareca. 4 im thick,andthe diameter ofthefibreluminais on average22 jim. Meanfibre is 1100 im.Ratherlongfibres> 1500pmoccurinLeptactinasenegambicaand length Pavettabrachycalyx Doricera (Tw 33222).ThejuvenilewoodsamplesofCaptaincookia margaretae, senniiandPavettacellulosashowrelatively shortfibres. trilocularis, Myonimaviolacea,Tennantia 120 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions IV. m . la-elem :Ilk wd ' Figure 19-27. Vessel pits viewedfromouterpit aperture:19, Pavettacellulosa, non-vestured pits; 20, Tennantiasennii,small unbranchedvesturesconcentratedaround outerpit aperture(vesturetype1) (bar = 1 pm); 21, Dictyandraarborescens(Tw 28571), vesturetype1; 22, Pavettanitidula,vesturetype1; 23, Tarennabipindensis,vesturetype1; 24, Tarennaattenuata, vesturesnotcompletelyfillingthepit chamber;25, Versteegiacauliflora,mostvesturesbranched,anastomosedand completely fillingthepithchamber(vesturetype2); 26, Captaincookiamargaretae,vesturetype2; 27, Doricera trilocularis(Tirvengadum 908), vesturetype2 (bar = 10 pm). This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions m :810 vesture bead-like vestures 28,Pachystylus type2; 29,Duperrea Figure28-37.Vesturing: pavettifolia, fillingthepit guelcherianus, vestures nearinnerpitapertures chamber;30,Leptactina arnoldiana, (bar= 10pm);31,Pachystylus present guelcherianus, at innerpitapertures andinnervesselwall;32,Myonima andvestured at innerpit violacea,vessel-ray vesturing pitselongated solomonensis 33,Captaincookia 34, Versteegia (MauriasiBSIP vestured; aperture; pitsoffibre-tracheids slightly margaretae, 35,Leptactina seretii (Louis1507),fibrepitspoorlyvestured 15875):fibrepitsvestured; (bar = 10pm);36,Pavetta brachycalyx vestured vestured. amoldiana, (Tw33222): innerpitapertures (bar = 10pm);37,Leptactina offibre simpleperforation This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions -Ii -- "$~w -m 0l .p, " ,-1 ne 71m- Figure38-46. Crystals:38 & 39, Tarennafusco-flava(Tw 41352), prismaticcrystalsin chamberedaxial parenchymacells (bar Jig.38 = 100 pm, bar fig. 39 = 50 pm); 40, Pavettanitidula,prismaticcrystalsin chamberedaxial parenchymacells (bar as in Jig.39); 41, Tarennafusco-flava(Tw 41352), chamberedaxial parenchymacells; 42 & 43, Nichallea soyauxii(Tw 32837), details ofprismaticcrystalsin chamberedaxial parenchymacells; 44 & 45, Captaincookiamargaretae,styloids(asterisks) presentin rows ofaxial parenchymacells; 46, Captaincookiamargaretae,elongate crystal. This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions mAlii n__r" A?Q .. IL 4:A .. .: m..... ...- .... Ai 'dlAk inaxialparenchyma 47, Ixoraferrea: (bar = 100pm);48, Leptactina present senegamelongatecrystals Figure47-53.Crystals: andchambered 49,Doriceratrilocularis bica,twoupright 908): druseinpith; (Tirvengadum crystals; raycellswith4 prismatic uniseriate sectionshowing 50 & 51,Tarennasupra-axillaris, rayswithsilicabodies(barfig.50 as infig.47); 52, tangential radialsectionwithdisjunctive Tarennaattenuata, (Tw33700),tangential raycellwalls(bar= 50 pm);53,Ixorabrachypoda vesselelements widevesselelements sectionshowing (arrows), (asterisks)withtwoperforations fibre(v),three fibriform andparenchyma cells(bar = 150pm). tracheids, 124 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions S. Jansen, P. De Block,H. Beeckman & E. Smets,Woodanatomy ofPavetteae (Rubiaceae) inLeptactina is scarcely seretii(Tw 34977,Tw 33276,fig.1). Diffuseaxial Axialparenchyma present is presentin Cladocerassubcapitatum, Dictyandraarborescens(fig.10), Duperrea parenchyma pavettifolia (fig.9), Ixorabrachypoda, I. sp.,all speciesofLeptactina (fig.2), Nichalleasoyauxii(Tw all speciesofPavetta(fig.3), Rutidea,Tarenna(fig.4; exceptT. 34928),Pachystylus guelcherianus, andTennantia sennii. supra-axillaris) occurspredominantly in all speciesofIxora (fig.5, 6), Axial parenchyma diffuse-in-aggregates Leptactinasenegambica,Myonimaviolacea, Tarennasupra-axillaris(fig. 12) and Versteegia solomonensis. inirregular, narrow bandsofthreecellswideis foundinwoodofCaptaincookia Axialparenchyma Doriceratrilocularis violacea,all speciesofVersteegia (fig.7), Myonima margaretae, (fig.8), andto a lesserextentinIxoralaxifloravar.laxiflora, L nigricans, andI. sp. I.ferrea(fig.6), I. graciliflora, whichis morethanthreeandsometimes morethan10 cells wide(ca. 100 plm) Axial parenchyma areirregular andnotfullycomposedof occursinDuperreapavettifolia (fig.9). Thebands,however, within becausevesselsandfibres arefrequently distributed thebands. axialparenchyma andcomposedofa low,2-to4-seriate orsquareraycells Theraysareheterocellular partofprocumbent cells. Exclusivelyuniseriate and long uniseriate marginsof upright raysoccurin Captaincookia seretii(Tw 34977,fig.18), Versteegia Leptactina margaretae, (fig.15),andTarennasupra-axillaris L L var. Ixora show 14), ferrea 50). laxiflora laxiflora,and I. graciliflora (fig. brachypoda, (fig. overa shortlength; thesemultiseriate uniseriate partsarenotmuchwider rayswhichmaybe 2-seriate arborescens thantheuniseriate (Tw 2416,fig. rayparts. Raysupto5 cellswidearefoundinDictyandra 13). cell walls are found.Disjunctive Perforated rayparenchyma raycells are simpleand frequently Ixora,Leptactina, Rutidea,and common, raycellsofe.g.Dictyandra, especiallyintheupright/square is usually> 1 mmandrayTarenna(fig.52). About12 ormorerayspermmarepresent. Rayheight areoften interconnected (Carlquist1988:202). VeryhighraysareobservedinRutideasmithii portions (Tw 40352). 1988: 192) occurinDuperreapavettifolia (fig.16): most typeI (Carlquist Raysofthepaedomorphic inthemultiseriate It is difficult cellsforma smallcluster andprocumbent portions. raycellsareupright sectionssincethelongmultiseriate to outlinetherayson tangential raycells marginswithupright cells. axialparenchyma resemble orintheraycells,largeprismatic intheaxialparenchyma smallrhombic crystalsoccurring Apartfrom in Tarennaeketensis, can be numerous axial cells. in are chambered parenchyma They crystals present T. Tw 39, (Tw 44603),T.petitii(Tw 35821, (Tw 32654, 41352;fig.38, 41), pavettoides T.fusco-flava inPavettacrassipes(Tw 17248),P. nitidula(fig.40), Nichalleasoyauxii Tw 38452), less numerous Thecrystals consist probably (Tw 34928,Tw 32837,Tw 33267;fig.42, 43) andHomollealongiflora. in structure as seen SEM have a somewhat laminated and of calcium oxalate (fig.43). The largely enclosed(fig.41chambered cellsshowno pitsandtheirshapeis similartotheoutlineofthecrystals chambers is oftenverylong,especiallyinTarenna(fig. ofthechainsofcrystalliferous 43). The length 38-39). arefoundinchambered senegambica(fig.48); note raycellsinLeptactina crystals Largeprismatic cells. in the axial shows no howeverthatthelatter parenchyma crystals species in Captaincookia margaretae(fig.44, 45), Ixora Styloidsbetween80-120gm longarepresent inlargeidioblasts ofaxialparenchymaandV.minor.Theyaresolitary Versteegia nigricans, cauliflora filltheidioblast andareusually The styloids inaxialdirection. tousoriginandareoriented completely aboveeach other(fig.44). Theyhaveelongatebodieswith foundin axial rowsof severalcrystals radiation of faces(fig.45). Undertheintense endsandtrapezoidal wedge-like tapered, asymmetrically tendtocrackuprapidly. thecrystals SEM microscopy, 125 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr. Pl. 68 (1999) of70-80pLm cellsin Captaincookia longarefoundinenlarged Elongatecrystals margaretae (fig. butconsiderably 46), Ixora ferrea(fig.47), andI. nigricans.Theyare similarin shapeto styloids shorter. We observeddrusesinthepithofDoriceratrilocularis 908; fig.49) andMyonima (Tirvengadum was also found in the of violacea.Thiscrystal pith Duperreapavettifolia. type Silicabodiesareabundantly intherayparenchyma cellsofTarennasupra-axillaris present (fig.50, 51). Chromeazurol-Stestforaluminium accumulation is negative forall woodsamplesinvestigated. Discussion studied showsexclusively orpredominantly Vessels.Woodofmostspecimens vessels,a feature solitary In contrast, thespecimen whichis commoninall Ixoroideae. andspecimen ofTarennasupra-axillaris Tw 23511 of Dictyandra arborescens and Tw 40352 of Rutidea smithiiare characterisedby the of2-4 vessels.Although 80-90%ofthevesselsare solitary in theother presenceofradialmultiples twospeciesstudied, theidentification ofTarennasupra-axillaris andRutidea specimensofthelatter smithii notoveremphasize thecharacter of (Tw40352)is beyonddoubt.One shouldtherefore grouping ofradialmultiples is possiblyaffected vesselsbecausea variablepercentage influenbyenvironmental ces.Tarennasupra-axillaris showsthehighest ofradialmultiples (ca. 80% ofvesselsoccur percentage inradialmultiples); thisspeciesis also unusualinothercharacters Koek-Noorman (see further). (1972) foundthatina specimenofTarennaincertaabout60% ofthevesselsaregroupedinradialmultiples. shenoticed thatthisspecimen waswrongly identified andis actually RandiawallichiiHook.f. However, oftheGardenieae wallichii(Hook.f.)Tirvengadum & Sastre],a tribeoftheIxoroi[nowTarennoidea occurinsomegenera, deae inwhichradialmultiples e.g.Amaioua,Duroia,Posoqueria,andRandia. Themorphology Vestures. anddistribution ofvestures intheintervessel examined pitswasthoroughly SEM. The two different of vestures described above are with the two informal using types congruent infratribal as 1 Bridson & Robbrecht in to occurs the related (1985): type groups proposedby genera whereas 2 in occurs the Ixora are and Tarenna, type group.Exceptions Ixora nigricans(type1) Pachystylusguelcherianus(type2). For Ixora nigricans,we shouldnotethattheaxial parenchymais morediffuse todiffuse-in-aggregates tothatoftheotherspeciesofIxora.Vestures oftypeI compared are identical tothevestures et described Jansen al. the for Coffeeae. As mentioned above, (1997b) by totheirtypeofvestured however,itis notalwayseasytoclassify specimens according pit.Ixora sp. showsvesturestransitional betweenthetwotypes.Moreover, within a singlespecimen, variation is Doricera for has vestured of of trilocularis, instance, possible. pits type2, butsomevestures frequently and morphological type I also occur.This mightin partbe explainedby thefactthatoccurrence invesselpitsvariesinmanycasesaccording variation ofvestures topittypesas demonstrated byOhtani & Ishida(1976).Itshouldbe noted,however, thatwe onlyfounddistinct variation between vesselpits and fibrepits.Clearly,duringSEM investigation, itis oftenverydifficult todetermine whichtypeof vesselpitis observed. The species withoutvestures(Pavetta cellulosa and Tarenna supra-axillaris) are remarkable because thecharacter ofvestured to be present theRubiaceae.Sincethe pitsis thought throughout ofthesetwospeciesis beyonddoubtwe wantto stressthattheirgenericpositionshouldbe identity reexamined. One maysuggestthatthelack of vesturing is apomorphic. Wood of Tarennasupraaxillaris showsseveralderivedcharacters butmorespecimens ofthisspeciesshouldbe examinedto draw a finalconclusion.Systematic literature on vestured thatthefeature is relatively pitsindicates wholefamilies andevenorders, conservative, & Smets1998), characterising (Jansen e.g. Gentianales 126 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions P. De Block,H. Beeckman & E. Smets, S. Jansen, Woodanatomy ofPavetteae (Rubiaceae) thatvestured andnon-vestured andMyrtales (VanVliet& Baas 1984).We believe,however, representhanhitherto known.Otherfamilieswithbothvesturedand nontativesco-occurmorefrequently vesturedtaxaareforinstance Loganiaceae(Mennega1980),Fabaceae(Quirk& Miller1983, 1985), (Miller1977,Gottwald1982),Oleaceae(Baas etal. 1988)andVerbenaceae(Mathew& Boraginaceae Shah 1983). tothedefinitions ofKoek-Noorman FibresinthePavetteaearefibre-tracheids (1969a) and according fibresare foundin Tarennaneurophylla thatseptatelibriform Baas (1986). It is remarkable and while libriform fibres are observed in seretii Tarenna (Tw 34977), non-septate Leptactina supraofthetwoTarennaspeciesis beyonddoubt,butwe couldnotconfirm axillaris.The identification the is also unusualcompared totheotherspeciofLeptactinaseretii(Tw 34977). Thisspecimen identity rather homocellular in itspossessionofuniseriate, nonmensofL. seretiistudied, raysandenlarged, borderedvessel-ray pits.ExceptforthegeneraAlibertiaand Posoqueria(Gardenieae) parenchyma libriform fibres(Koek-Noorman whichhavenon-septate to 1972),woodofall Ixoroideaeis suggested etal. 1997b). havefibre-tracheids (Jansen ThemostcommontypesintheIxoroideaeandPavetteae arediffuse anddiffuse-inAxialparenchyma. for instance in and These are Tarenna. Short Pavetta, Rutidea, Leptactina, aggregates. types present are characteristic for and bands Doricera, Captaincookia, Myonima, Versteegia. parenchyma Generally, tobe fromdiffuse theevolutionary (themost parenchyma typesis concluded sequenceofapotracheal to transitional bands bands are (wide parenchyma through types(diffuse-in-aggregates), primitive type), means of mosthighly 1937, (Kribs by increasing parenchyma aggregation Carlquist1988). specialised) inthisstudy.Specimens inthewoodexamined ofIxorashowaxialparenchyma isevident Thistendency to smalland narrowparenchyma or diffuse-in-aggregates bands(especiallyI. fromdiffuse, varying link the with the Ixora forms a between diffuse In this andthe context, genus ferrea). genera parenchyma Doricera and Other wood anatomists derived)generaCaptaincookia, Versteegia, Myonima. (probably in broken, linesor finebandsin ixora. suchas Williams(1936) foundaxialparenchyma tangential and in Versteegia bandsand Ridsdaleetal. (1972) notedthattheparenchyma (short longparenchyma to with that in Alibertia and is Duroia (bothbelonging Gardenieae-Gardeniinae). islands) comparable in Duperreapavettifolia is ratherunusualforrubiaceouswood: it consistsof Axial parenchyma bandsas wellas diffuse wideparenchyma parenchyma. irregular, is fromtheheterogeneous inthewoodraysofdicotyledons typeI to the Rays. The generaltendency than with more ones 1935, II, (Kribs primitive heterogeneous rays homogeneous homogeneous type uniseriate and multiseriate are the both Within Pavetteae, 1988). rays present. Predominantly Carlquist thewoodsamplesoftheIxora andVersteegia. uniseriate raysoccurinIxora,Captaincookia, Although by otherinvestigators. groupexaminedin thisstudyare oftenjuvenile,ourresultsare confirmed of cells are described Ridsdale et al. (1972) in uniseriate by rayscomposed upright Predominantly and V. solomonensis. Koek-Noorman also found uniseriate (1972) Versteegiacauliflora exclusively over a short distance in Ixora biseriate). (or rarely very rays Ridsdaleetal. (1972) forinstance raycellwallsarecommonintheIxoroideae. reported Disjunctive cellsofVersteegia. intherowsofprocumbent andupright elements manydisjunctive areabsentinthesecondary inclusions Mineralinclusions. crystals,mineral Apartfromsmallrhombic Pavetta ofthegeneraCladoceras, Dictyandra, Leptactina (exceptL. senegambica), xylem Pachystylus, occur Numerous Rutidea,andTennantia. crystals largeprismatic (exceptP. crassipesandP. nitidula), inNichalleasoyauxii(Tw 34928, inLeptactina inmostoftheTarennaspeciesstudied, senegambica, inPavettacrassipes(Tw 17248)andP. nitidula. These Tw 32837,Tw 33267),andto a lesserextent intheraycellsas demonstrated forthetribeCoffeeae(Jansen etal. arenotabundantly present crystals 127 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr. Pl. 68 (1999) axialparenchyma cells.ThewoodsampleofLeptactina 1997b),butoccurinchambered senegambica notconfirmed) studied(voucherspecimennotseen;identity is unusualbecausethecrystals occuronly Table3. Diagnostic woodanatomical ofthegenera characters studied. -= absent, + = present, D = diffuse, RM= radial S = solitary, DA= diffuse-in-aggregates, NB= narrow bands, multiples, inchambered WB= widebands, CAP= prismatic axialparenchyma, CRP= prismatic inchambered crystals crystals ray ST= styloids, EC= elongate SB = silicabodies parenchyma, crystals, mineral inclusions genus vessel ray vesturing septate axial fibres parenchyma width grouping type Captaincookia S 2 - NB 1 Cladoceras S 1 - D 2-4 Dictyandra S (RM) 1 - D 2-5 Doricera S 2 - NB 2-4 Duperrea S 2 - D-WB 2-4 Homollea S ? - D 2-4 CAP Ixora S 1-2 - D-DA-NB 1-2 EC-(ST) D-DA 2-4(1) CRP Leptactina S 1 - (+) Myonima S 2 - DA-NB 2-4 Nichallea S 1 - D 2-4 Pachystylus S 2 - D 2-4 Pavetta S 1 - D 2-4 Rutidea S (RM) 1 - D 2-4 Tarenna S (RM) 1 - (+) D (DA) 2-4 (1) Tennantia S 1 - D 2-4 S 2 - DA-NB 1 Versteegia ... L.. ST-EC CAP CAP CAP (SB) ST .. ...... in chambered in chambered cellsareveryrarein thesecondary crystals raycells.Prismatic xylemof in some speciesof Coutarea(Portlandiagroup)and Pavetta Rubiaceae and wereonlyreported (Metcalfe& Chalk1950:769). arealso observed inthesecondary Itis worth thatprismatic phloemofLeptactimentioning crystals andL. senegambica, na seretii(Tw 33276),L. pynaertii, as wellas inthepithcellsinL. mannii.The inthebarkorpithis beyongthescopeofthiswoodanatomical occurrence ofcrystals butitwould study, toincludeobservations onthesefeatures forfuture anatomical be interesting research ofthePavetteae. andIxora.These Captaincookia StyloidsorelongatecrystalsarefoundinthegeneraVersteegia, rare in wood. of in are are rubiaceous Previous the reports styloids family fromJanssonius crystals andIxora),Koek-Noorman (1926,p. 200),Chattaway (1955;Hymenodictyon (1972;Ixora grandiflora and andI. macrophylla), Ridsdaleetal. (1972; Versteegia Richter & Schmitt (1987). The cauliflora), of in Standl. latterauthors the (inc.sed.). spectabilis reported presence "mega-styloids"Cosmocalyx inthebarkofIxoraandMyonima. Beille(1947) and Solereder(1893) notedthepresenceofstyloids inthesecondary Doriceraand largestyloids phloemofIxorasp.Although Chang(1951) also reported intheirwood,thiscrystal a valid orelongatecrystals constitutes Myonimado notshowstyloids type 128 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions S. Jansen, P. De Block,H. Beeckman & E. Smets,Woodanatomy ofPavetteae (Rubiaceae) taxonomic feature thatcan be employed tocharacterise theIxoragroupandseparateitfromtheother Pavetteae. Itespeciallyconfirms thecloserelationship betweenCaptaincookia, IxoraandVersteegia. Druses in thepithofDoriceraandMyonima thecloserelationship betweenboth possiblysupport thepithofmostgenerawasnotexamined andtherefore taxonomic conclusions However, should genera. be avoided.Druseswerealsoreported (1893) inthebarkofIxoraandMyonima. bySolereder Similarly, thepresence ofidioblasts a singledrusewasdemonstrated containing byTange(1996a,b) inthepithof Aleisanthia andAleisanthiopsis (Rondeletieae). Silica bodies are veryrarein woodoftheRubiaceae:theyhaveonlybeenreported in woodof etal. 1995). Therefore, Nauclea,andHallea (Carlquist1988,TerWelle1976,Vrydaghs Mitragyna, their inTarennasupra-axillaris occurrence is remarkable. ThisMadagascanspeciesdiffers further from theotherTarennaspeciesstudiedbythepresenceofradialvesselmultiples, uniseriate raysandaxial inaggregates. parenchyma inthesecondary Environmental influences arevisibleinthepresenceofgrowth xylemofthePavetteae in and differences in certain suchas vesseldiameter, vessel characters, rings quantitative occurring and fibre wall thickness in the lianescent oflianasin frequency genusRutidea).Specificfeatures (e.g. thisgenusare:twodifferent sizesofvessels,a rather andthinfibrewalls.Koeklargevesseldiameter than Noorman vessels more 100 wide in Rutideamembranacea; thevesselsinour (1972)reported ipm 8 mm)ofR. membranacea, areon average30 pmwide.The however, juvenilewoodsample(diameter lianas Tarennafusco-flava(Tw 32654,Tw 41352) andT. eketensis also havetwodistinct sizes of incomparison vesselswiththelatter widerandmorecircular tothoseofotherTarennaspecies. Relationshipsand evolution ComparisonwithothertribesoftheIxoroideae Wood ofthePavetteaeagreeswiththesecondary features: xylemofotherIxoroideaein thefollowing cell occurrence of and vessels, fibre-tracheids, walls, disjunctive rayparenchyma solitary frequent Thetribediffers absenceofhighaluminium accumulation. woodanatomically fromtheCoffeeaebythe intheraycells(Jansen absenceofprismatic etal. 1997b).Prismatic arealso conspicucrystals crystals the tribes and some in cells of Aulacocalyceae, Octotropideae generaoftheGardenieae ouslypresent ray theIxoroideaein aretheonlytribewithin On theotherhand,thePavetteae observation). (S.J.:personal andcrystals inchambered axialparenchyma cellsareknown.Other whichstyloids orelongatecrystals withinthesubfamily are thedistribution whichshowsome variation of wood anatomicalfeatures and theraystructure. Afterpreliminary observations on thevariationof apotrachealparenchyma we believethatvariation andOctotropideae, investure vestured Aulacocalyceae pitsintheGardenieae, tothetwovesture above. is restricted typesdescribed morphology Ixora fromthe Koek-Noorman(1969b, 1972) concludedthatit is notpossibleto distinguish andstatedthatthewoodoftheGardenieae and"Ixoreae"is Gardenieaeon thebasisofwoodanatomy andtribalcircumscriptions theclassification havechangedthoroughly verysimilar.Although system dataagreewithherresultsandindicate thattheIxoroideaeare sinceherstudies,ourwoodanatomical is foundinthe Within thissubfamily, mostwood anatomicalvariation woodanatomically homogeneous. theneotropical Pavetteaeand,toa lesserextent, Gardenieae-Gardeniinae. The Ixora group indicate a close resultsofthepresent One ofthemostinteresting studyis thefactthatwoodcharacters andVersteegia. Thesegeneraare thegenera between Myonima affinity Ixora,Doricera,Captaincookia, in features: bands(all genera,butnotconsistently characterized present parenchyma bythefollowing drusesinthepith(Doriceand/or Ixora,Versteegia), Ixora),styloids (Captaincookia, elongatecrystals 129 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr. Pl. 68 (1999) andto a lesserextent ra, Myonima),vesturetype2 anduniseriate rays(Captaincookia,Versteegia a of wood anatomical characters see table 3). Ixora)(for summary diagnostic Otherauthorsalreadyexpressedtheidea of a verycloselyrelated"Ixora group."Bridson& Robbrecht differences between Ixora (1985) intheirreviewofthetribewereawareofthefundamental andtheothergeneraofthePavetteae:theyrecognized and its close relatives, twoinformal groups, notablygenerarelatedto Ixora and generarelatedto Tarenna.De Block (1997), afterreviewing inthePavetteae, madea preliminary andbiologicalcharacters anatomical, palynological morphological, the and confirmed close between the cladistic five of the analysis genera relationship Ixoragroup.Some evengo further; a cladisticanalysisoftheIxoroideae basedona combination ofmolecular authors and & Bremer1996;Andreasen datawasrecently (Andreasen 1997)andindicated presented morphological thatthetribe PavetteaesensuBridson& Robbrecht is paraphyletic. toAndreasen (1997) the According generaIxora, Doricera, Captaincookia,Myonimaand Versteegiashouldbe excludedfromthe Pavetteaes.s. as a tribeIxoreae(see table2). Considering thehomogeneous secondary xylemofthe woodanatomy seemstojustify theacceptanceofa separatetribeIxoreaesensu Ixoroideae, subfamily Ifnot,thewoodanatomical stricto. distinction theIxoragroupandtheotherPavetteaeshould between be emphasized andtheIxoragroupshouldbe givenatleastsubtribal rankwithin thePavetteae. The positionofDuperrea WoodofDuperreapavettifolia differs fromtheother Pavetteae bands, bytheunusualwideparenchyma theraycomposition andthebead-likevestures. Thiscorroborates theexclusionofthegenusfromthe as alreadysuggested tribePavetteae, (1997a) andAndreasen (1997). The byDe Block& Robbrecht former examinedmorphological, anatomical authors andpalynological characters oftheIndochinese that itshouldbe removed fromthePavetteae andtransferred toeither the genusDuperreaandconcluded tribeGardenieae orAulacocalyceae. Theirreasoning wasbasedprimarily ontheabsenceofan adaxial a reduction oftheseed-coat, andthecomplete oftheovulesin placental tissueat excavation, embedding theflowering characters which are for these two tribes. to Andreasen stage, typical According (op. cit.) tothetribeGardenieae. Woodanatomical characters seemtocorroboDuperreashouldbe transferred ratethissince(wide)parenchyma bandsarerareinthesubfamily Ixoroideaeandonlyreported inthe Ibetralia and all to Alibertia, Duroia, genera Borojoa, (Alibertia?), Kutchubaea, belonging the Gardenieae(Koek-Noorman 1969b,1972). Otherrelationships Interandinfrageneric within thePavetteae arestillunclear. relationships ExceptforthegenusIxoraand itscloserelatives considered (see above),oneothergroupofgenerais generally closelyrelated, notably thegeneraLeptactina, andColeactina.Thesethreearecharacterized others, Dictyandra by,amongst ovulesandwiththeareaofattachment largeU-shapedplacentaswithnumerous coveringthewhole characters whichareconsidered tobe primitive intheRubiaceae(De Block& lengthoftheplacenta, Robbrecht1997b;De Block 1997). Woodanatomy ofLeptactinaandDictyandra lacksthefeatures characteristic fortheIxoragroup,butitprovides nofurther toconcludethatthetwogenera information are moreprimitive or morecloselyrelatedto each otheras forinstanceto Pavettaor Nichallea. Leptactinaseretii(Tw 34977) is a specieswhichshowsderivedcharacters, notably septatelibriform fibresanduniseriate, nearlyhomocellular rays. The twogeneraRutideaandNichalleaarealso considered tobe closelyrelated(Bridson& Robbrecht1985;De Block 1995).No woodanatomical arecorroborating features this. within Thelargest unresolved thetribePavetteae is thedelimitation ofthegenusTarenna. problem Variablecharacters aree.g.placentation, ovulenumber, rumination oftheendosperm andthestructure oftheseed-coat.Bridson(1979) questioned whether African andnon-African speciesofTarennatruly sherecognized six informal belongto thesamegenus.Furthermore, groupsfortheAfromadagascan ofTarennas.l.In thisstudy, material oftwoofthesegroups[notably ofBrison'sgroup representatives 130 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions S. Jansen, P. De Block,H. Beeckman & E. Smets,Woodanatomy ofPavetteae (Rubiaceae) I andIV] was examined withoneAsiaticspecies.Thetaxaarelistedintable4 withreference together tothepresenceofcrystals inchambered axialparenchyma cells.Itis interesting tonotethatall species withcrystalliferous chambersbelongto Bridson'sgroupI. ExceptforTarennabipindensis and T. all from I show this feature. Other wood anatomical differences precidantenna, species group crystal cannotbe found.It is clearthattheoccurrence ofprismatic in thegenusTarennas.l. needs crystals inall ofBridson'sgroups. further verification Table 4. The Tarennaspeciesstudiedand thepresenceofprismatic cells. crystalsin chamberedaxial parenchyma toBridson's(1979)informal classification. Speciesarranged according groupI Icrystals absent T.bipindensis T eketensis Tw38302 Tw35792 present T fusco-flava T fusco-flava Tw32654 present Tw41352 present T pavettoides Tw44603 present T petitii T petitii Tw35821 present Tw38452 T precidantenna Tw38479 present absent groupIV T graveolens Tw39207 absent T neurophylla Tw45140 absent T supra-axillaris Tw44778 absent(presence ofsilicabodies) Asiaticspecies T attenuata Tw42126 1absent Bridson's Furthermore, T. neurophylgroupIV,ofwhichthreespecieswerestudied(T. graveolens, T. is woodanatomically and The AfricanT. neurophylla shows la supra-axillaris), heterogeneous. the libriform is remarkable the fibres; MadagascanspeciesT. supra-axillaris septate by presenceof radialvesselmultiples, silicabodies,uniseriate axial in small lines and thepresenceof rays, parenchyma libriform fibres. Wood demonstrates that more taxonomic research on Tarenna, anatomy non-septate is required. especiallyofMadagascanrepresentatives Evolutionarytrends We suggestthefollowing trends within thesecondary xylemofthePavetteae: specialisation ofapotracheal axialparenchyma fromdiffuse tobanded. (1) Aggregation bordersto muchreducedor (2) Raysfrom2-3-4-seriateto uniseriate; vessel-ray pitsfromdistinct borders. simple to(non-)septate libriform fibres. (3) Fibresfromfibre-tracheids tointerpret thecorrelation ofthetwotypes However, (4) Vesturetypesaredifficult phylogenetically. describedinthePavetteaewithotherwoodanatomical features allowsthephylogenetic interpretation thanthoseoftype2. In otherwordswe proposea trend thatvestures oftype1 aremoreprimitive from vestures neartheouterpitaperture to largervestures a network that forming verysmall,unbranched NotethatinVan Vliet'sspecialisation seriesofvesture occludesthepitaperture. typesintheMyrtales becomemoreconcentrated aroundthe (Van Vliet& Baas 1984;typesA, B 1,B2, andB3) thevestures trunk-like bases. and pitapertures develop 131 This content downloaded from 193.190.117.53 on Tue, 26 Mar 2013 05:06:54 AM All use subject to JSTOR Terms and Conditions Syst.Geogr.Pl. 68 (1999) andstyloids. thesecrystals, itis verydifficult topicture ofelongate (5) Development crystals Apartfrom as or theothercrystal types primitive specialised. - We thankDr. J.Koek-Noorman andProf.Dr. E. Robbrecht forhelpful comAcknowledgements. of K, L andP areacknowledged fortheloanofherbarium Theherbarium curators material and ments. forthepermission toexecuteanatomical studies.StevenJansen holdsa bursary oftheResearchCouncil of theK.U. 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