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Ixoroideae Rubiaceae Grana39: 90± 102, 2000 Pollenmorphological variation in Vanguerieae (Ixoroideae ± Rubiaceae) FREDERIC LENS,STEVEN JANSEN,SUZY HUYSMANS, ELMAR ROBBRECHT andERIK SMETS Lens, F., Jansen, S.,Huysmans,S .,Robbrecht,E . & Smets, E. 2000. Pollenmorphological variation in Vanguerieae ( Ixoroideae± Rubiaceae ) . ±Grana39: 90± 102 . ISSN0017-3134 . TheVanguerieae is alargetribe of the subfamily Ixoroideae ( Rubiaceae) andconsists ofabout 500 speciesin 27 genera . This studygives a detailedpollen morphological description of 30species from 16 genera,based on light microscopy and scanning electron microscopy . TheVanguerieae are considered tobe stenopalynous, although there is someevidence to question this . First, thereis adifferencein sexinetypes that clearly de® nes Keetia and Psydrax ( bothreticulate ) , and Canthium and Vanguerieae ( bothperforate with verysmall perforations ) fromthe other Vanguerieae investigated . Second,pollen grainscorroborate the subgeneric delimitations of Canthium sensuBridson . Pollenmorphologically the tribeis characterisedby ( 1) theshape of theectoapertures which is intermediatebetween a colpusand a porus, and ( 2) thepresence of a costaoriented perpendicular to the ectoaperture and has a formwhich is correlatedwith theshape of the costa . FredericLens, Steven Jansen, Suzy Huysmans & Erik Smets,Laboratory of PlantSystematics, Institute ofBotany and Microbiology, K .U.Leuven,Kardinaal Mercierlaan 92, B-3001 Leuven; Elmar Robbrecht, NationalBotanic Garden of Belgium, Domein van Bouchout, B-1860 Meise; Belgium . E-mail: [email protected]. ( Manuscriptaccepted 29 September 2000 ) Withnearly 500 species arranged in 27 genera, the blanceswould most probably reduce the number of genera . Vanguerieaeis one of the larger tribes of the subfamily Afterthe revision of Canthium by Bullock ( 1932) , Bridson Ixoroideae ( Rubiaceae) . Itis widespread in tropical Africa, ( 1985,1986, 1987 a, b, 1992) refutedthe opinion of Asia,Australia and the isles of the Paci® c Ocean,but the Verdcourt. Sheaccepted and broadened, on the basis of largestnumber of species occurs in Africa and Madagascar morphologicalfeatures, several genera of Canthium s.l., ( Bridson1987 a) . Thehabit is generally shrubby, but also e.g. Psydrax, Pyrostria, Keetia and Multidentia . Further- fewlarge trees and lianas occur . Thetribe combines many more,she recognised four subgenera for the remaining distinctivefeatures, e .g. axillarypaired in¯ orescences, species ( ~ Canthium s.s.) , namely Canthium, Afrocanthium , valvateaestivation, secondary pollen presentation, one Lycioserissa and Bullockia. Thesetaxonomic changes are pendulousovule per locule, pyrenes with an apical alsosupported by otherauthors, such as Tilneyet al . ( 1988) germinationslit, and seeds with soft oily endosperm and whostudied anatomical features of young stems of South verylarge embryos ( Robbrecht1988 ) . Africanspecies of Canthium s.l. Accordingto the classi® cation of Schumann (1891) , the Themacromorphological homogeneity of the tribe becomes Vanguerieaewere placed in the supertribe ` Guettardinae’ alsoevident in the pollen morphology . Robyns ( 1928:6) togetherwith the Alberteae, Guettardeae, Retiniphylleae and mentionedthat pollen grains in the Vanguerieae are `tre Ás Knoxieae. Thisphylogenetic relationship was adopted by semblablesdans tout le groupe’ . Verdcourt ( 1958) came to the Robbrecht ( 1988, 1994) whounited these tribes in his sameconclusion and described the grains as 3-porate with an subfamilyAntirheoideae . Moleculardata have demonstrated unsculpturedsexine . However,Robbrecht ( 1980) found morerecently that the subfamily Antirheoideae is polyphyletic severalsexine types within the tribe . Moreover,Tilney & andthat the Vanguerieae and the closely related Alberteae van Wyk ( 1997) foundpollen morphological support for the belongto the Ixoroideae ( Bremer et al. 1995,Andreasen & division of Canthium s.s. intothree subgenera . Otherstudies Bremer1996, Bremer 1996, Bremer & Thulin1998, alsoreveal pollen morphological variation within the Andersson& Rova1999, Andreasen et al . 1999,Rova 1999 ) . Vanguerieae,although the tribe is still considered to be Althoughthe Vanguerieae are well characterisedmor- stenopalynous ( Verdcourt1987, Igersheim 1989 ) . phologically,the generic delimitation remains problematic . Theinside ornamentation of pollengrains in Rubiaceae is Robyns ( 1928) wasthe ® rsttaxonomist who dealt with this knownto have systematic value, as ® rstmentioned by problem. Herevised the tribe, without treating Canthium, Frenchresearchers ( e.g. Malplanche1971, Lobreau-Callen whichis the largest genus that most likely plays a keyrole in 1978,Van Campo 1978, Keddam-Malplanche 1980 ) . Since thetribe’ sphylogeny ( Bridson1992 ) . Thehigh degree of Huysmanset al . ( 1994) describeda newmethod for breaking homogeneitywithin the Vanguerieae is discussed by pollengrains with glass beads, observations of the inside of Verdcourt ( 1958,1987:123 ) whocommented on `the thegrains became more accessible ( e.g. Jansen et al. 1996, poornessof characters used for generic delimitation’ . DeBlock & Robbrecht1998, Bosser & Lobreau-Callen Accordingto Verdcourt, the strong morphological resem- 1998,Huysmans 1998 ) . Newfeatures, such as different Grana 39 ( 2000) # 2000Taylor & Francis . ISSN0017-3134 Pollenmorphology of Vanguerieae 91 ornamentationtypes of the inner nexine and the large MATERIAL ANDMETHODS diversityof endoapertures, have shown their taxonomic importance. Thisis demonstrated for example in the Pollenof 30speciesout of 16 genera were removed from herbarium Coptosapelteae ( Huysmanset al . 1993, 1994) and in the specimensat the National Botanic Garden of Belgium ( BR) and Isertieae s.l. ofwhich characters from the inside of the wereacetolysed following the method of Reitsma ( 1969) . The pollenwere included for the ® rsttime in a cladisticanalysis specimensinvestigated are listed separately at the end of this paper . ( Huysmanset al . 1998) . InVanguerieae, however, inside LMpreparationswere made in glycerine jelly . For SEM observa- observationsof pollen grains were only made for the small tions, pollengrains were mounted on an aluminium stub, coated with goldand observed with aJEOLJSM-5800 LV or JSM-6400 . genus Tapiphyllum ( Havard& Verdcourt1987 ) and few Pollengrains were broken using glass beads ( Huysmanset al . 1994) . otherrepresentatives of the tribe ( Igersheim1989 ) Thepolar axis and equatorial diameter were measured on ten Thepresent study gives a detailedpollen morphological grainsin equatorial view with LMusinga cameralucida . Other descriptionof selected species of the Vanguerieae, including measurementswere made on SEM-micrographs . observationsof the inside ornamentation of the pollen Terminologyfollows theGlossary of Pollenand Spore Terminol- grains. ogy ( Punt et al. 1994) . Theterminology of pollen shape in polar Figs. 1 ± 6. Pollenin equatorial view . SEM. ( 1) Cuvieralongi¯ ora ,perforatespheroidal grain with congruentapertures; ( 2) Canthium parasiebenlistii ,suboblateshape, microverrucae ( detailin Fig . 28) inbands around psilate apoporium; ( 3) Pyrostriabibracteata , subo- blateperforate grain; ( 4) Psydraxhorizontalis ,suboblatereticulate grain; ( 5) Keetiahispida , idem; ( 6) Psydraxsubcordata var. subcor- data,spheroidalreticulate pollen . Grana 39 ( 2000) 92 F. Lens et al. Figs. 7 ± 14. Pollenin polar view . SEM. ( 7) Canthiumseti¯ orum ,ambview circular, perforated sexine; ( 8) Pyrostriabibracteata , amb viewcircular, perforate tectum; ( 9) Canthiumparasiebenlistii ;polarview circular, granules ( detailin Fig . 28) inbands around psilate apoporium; ( 10) Pygmaeothamnuszeyheri ,circularamb view, perforate sexine with psilateapoporium; ( 11) Lagyniaspallidi¯ ora , amb viewnearly circular, perforate sexine; ( 12) Fadogiaschmitzii ,polarview triangular with convexsides, perforatetectum; ( 13) Tapiphyllum cinerascens var. laetum, idem; ( 14) Psydraxhorizontalis ,ambview triangular with convexsides, reticulatesexine . Grana 39 ( 2000) Pollenmorphology of Vanguerieae 93 Grana 39 ( 2000) 94 F. Lens et al. viewfollows Reitsma ( 1970) . Terms for shapeclasses in equatorial instance Canthiumlactescens ( Fig. 25) and Psydraxsub- vieware adopted from Erdtman ( 1971) . cordata var. subcordata ( Fig. 26) . Ectocolpiwithin the Themeasurements are always mean values . Asummaryof the Vanguerieaeare scarcely present and always short with pollenmorphological features per species is listedin Table I . obtuse ends ( e.g. Vangueriainfausta ) oracute ends, such as in Fadogiastigmatoloba ( Fig. 15) . Themargin of the RESULTS ectoapertureis often obviously demarcated, except in Canthiumseti¯ orum ( Fig. 21) and Keetiagueinzii ( Fig. 22) ; Size sometimesthe ectoaperture has irregular margins because of Pollengrains are generally medium sized ( P: 44.9 mm and E: sexinebulges, especially in Canthiumlactescens ( Fig. 25) and 48.9 mm) . Thelargest pollen grains are observed in Cuviera Canthiummundianum . Thelargest ectoapertures are nigrescens ( P: 65.1 mm, E: 67.1 mm) ,thesmallest grains in observed in Cuvieranigrescens , Pachystigmapygmaeum , Canthiummundianum and C. parasiebenlistii ( P: ca 29 mm, E: Lagyniaspallidi¯ ora and Psydraxhorizontalis ( length ca ca 32.5 mm) . 16.5 mm;width ca 10 .5 mm) ;thesmallest ones in Canthium parasiebenlistii and Canthiummundianum ( length ca 5 mm; width ca 4 mm) . Ectoaperturemembranes are usually Shape coveredwith few microverrucae ( e.g. Fig. 15) . However, in species of Rytigynia and Vangueriopsis thesestructures
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