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Evolutionary biology of Brunellia (Brunelliaceae, Oxalidales)
Orozco Pardo, I.C.
Publication date 2001
Link to publication
Citation for published version (APA): Orozco Pardo, I. C. (2001). Evolutionary biology of Brunellia (Brunelliaceae, Oxalidales). Universiteit van Amsterdam.
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Download date:27 Sep 2021 POLLENN MORPHOLOGY OF BRUNELLIA RUIZ & PAVÓN AND RELATEDD TAXA1
Chapterr 6
Claraa InésOrozco
ABSTRACT T Pollenn grains of 24 species of Bruneliia were examined with LM and SEM in order to find additional characterss for phylogenetic analysis. The pollen grains were found to be 3- colporate, tectate and to have aa variable ornamentation which forms an almost perfect continuum. Five categories of exine ornamentation weree observed: striate reticulate (large lumina and high muri), finely reticulate, modified reticulate (muri andd lumina irregular in shape and at various levels), modified rugulate (irregular and protruding tectal elements)) and punctate (the lumina are smaller and round to slit-shaped). The exine ornamentation providess phylogenetic information for some groups of species, and in some cases it is correlated with
vegetative,, inflorescence and fruit characteristics. The punctate type could be the plesiomorphic character
state.. Relationships in the pollen morphology of Brvneilia and certain genera of Cunoniaceae are discussed.
RESUMEN N Ell polen de 24 especies de Bruneliia fue examinado en el microscopio de luz (ML) y el microscopio
electronicc (MES) para encontrar caracteres adicionales y usar esta informacion en el estudio de relaciones
filogenéticas.. El grano de polen es 3-colporado, tectado y con una alta variabilidad en la ornamentacion
dee la exina la cual forma un casi perfecto continuo. Se observaron cinco categories en la ornamentacion
dee la exina del grano de pollen:reticulado a finamente reticulado, reticulado modificado (muros y luminas
dee forma irregular y en varios niveles), rugulada modificada (irregular y proyectando elemento&tectales) y
laa ornamentacion de tipo punteado (las lüminas son pequeöas redondeadas o en forma de lineas). La
ornamentacionn de la exina tiene importancia filogenética para algunos grupos de especies y presenta
correlacionn con algunos caracteres vegetativos de inflorescencias y del fruto. El tipo de ornamentacion
punteadoo parece ser e! estado plesiomórfico. Se discuten las relaciones del grano de polen de Bruneliia
yy ciertos géneros de Cunoniaceae.
»:cniminGm"» » CHAPTERCHAPTER 6 POLLEN MORPHOLOGY OF BRUNELUA RUtZ & PAVON AMD RELATED TAXA
91 91
6.11 INTRODUCTION
Thee genus Bruneilia Ruiz & Pavón includes 54 species (Orozco in prep.) and is currently considered thee only genus of the family Brunelliaceae Bruneilia is neotropical and widefy distributed in Andean and subandeann forests where high humidity and frequent cloudiness prevail Species of Bruneilia also occur fromm Mexico to Bolivia and in the Greater Antilles The highest number of species. 34 is present in
Colombiaa (Chapter 1)
Mostt of the earlier taxonomie classifications indicated relationships between Brunelliaceae and
CunoniaceaeCunoniaceae and often both families were placed in the order Rosales (CRONQUIST 1981, DAHLGRËN 1980.
TAKHTAJANN 1981.THORNE 1983). CUATRECASAS (1970) also noted a close relationship between Bruneilia and
Cunoniaceaee especially with the genera Spiraeanthemum and Acsmithia based on vegetative and anatomicall characters The relationship of the two families, however, is currently under discussion; both familiess have been included in the order Oxalidales ( APG 1998). Research on the position of Bruneilia hass also been earned out by HUFFORD and DICKISON (1992) who placed Bruneilia in the family of Cunoniaceae. whereass OROZCO (1997) suggested that Bruneilia, Spiraeanthemum and Acsmithia form a separate family, cioselyy related to the Cunoniaceae. However, BRADFORD ( pers comm ) from his studies on molecular data, considerss that Brunelliaceae must be retained as a monotypic family He indicates that Brunelliaceae showw close relationships with Cephalotus (Cephalotaceae) and with Cunoniaceae. but Brunelliaceae probably doess not fall within the Cunoniaceae clade It was indicated by HICKE^ and TAYLOR (1991) and also by
OROZCOO (1997) that Cunoniaceae is not a monophyietic group
Cunoniaceaee Brunelliaceae Eucryphia (Eucryphiaceae) and Davidsonia (Davidsoniaceae) are consideredd by Dcosc. (1989) to comprise the basal group in the rosalean complex Connaraceae is anotherr closely related family to Brunelliaceae It used to be included m Rosales but now it is placed togetherr with Brunelliaceae and Cunoniaceae in the order Oxalidales (AGP 1998) Most of the genera of
Connaraceae,, with exception of Agelaea. Cnestidium and Cnestis share the presence of five vascular tracess in the carpels with Bruneilia. Spiraeanthemum and Acsmithia (DICKISON 1971) Other characteristics sharedd among Bruneilia. Spiraeanthemum and Acsmithia and also by Connarus, include the absence of petalss and the apocarpous gynoecium. The epitropous ovule and the absence of a spectahzed xylem with scatariformm pitting are aiso seen in Bruneilia. Spiraeanthemum and Acsmithia (DICKISON 1975. 1980,
CUATRECASASS 1970. 1985). From molecular data it appears (BRADFORD pers comm.) that Brunelliaceae is closerr to Cunoniaceae and Cephalotaceae than to Connaraceae among the families of the order Oxalidales CHAPTERSCHAPTERS POLLEN MORPHOLOGY OF BRUNELUA RUIZ &PAVÖN AND RELATED 'AX A
92 92
Hu^c=:: and 0-->-5QN (1992), OROZCC (1997) and BR~:^O&O (pers comm i consider the genera Eucryphta andd Davidsonta within Cunomaceae
Thee first observations of pollen grams of Brunellia were made m S comociadifolia by E=D_V-N
(1952)) who with regard to sexine described the grams as having a rough surface and somewhat irregular sculpturee Basak (1967) described the pollen gram of B boliviana and B comociadifolia He noticed differencess in the surface pattern of the two species in B boliviana an echinuiate reticulate pattern being observed,, whereas the pattern in S comociadifolia was seen as irregular to slightly echinuiate Fossil
pollenn of Brunellia was first reported by GRAHAM and JARZEN (1969) from Puerto Rico They did not find differencess between the fossil grams of Brunellia and the modem species including B. comociadifolia
SALOMONSS (1986) described fossil pollen of B goudotii using LM He observed intectate or semitectate
pollenn grains with verrucate, fossulate, scabrate or microreticulate sculptures MAF nc OREN A (1970} studied
severall species of Brunellia using LM. He found two basic patterns of exme structure, one with a finely
reticulatee sexine without differentiated columellae and the other intectate with verrucose processes Results
reportedd by OROZCO (1991) differ from those of Marticorena as the former did not find intectate pollen
gramss in the SEM observations
Theree are few complete studies on the pollen morphology of groups considered close to Brunellia
HiDELxx and FERGUSON (1976) studied pollen grains, under light (LM) and scanning electron microscopy
(SEM)) of Saxifragaceae sensu lato, including Saxifragaceae sensu stncto. Cunomaceae Escailontaceae
andd Hydrangeaceae Pollen morphology in Connaraceae was studied by DIC SEM M Observationss on pollen grams of groups related to Brunellia, such as those of WAL (1975).. indicate that tncolporate grains and finely reticulate (small luminaj sculpture are especially common inn many basal groups of Rosidae It seems that the oblate triangular grains with apertures at the angles (angulaperturate)) is a character present in the basal groups of Rosidae as it is present in Cunomaceae Thiss character is also present m Connaraceae, however this family was not considered a basal group of Rosidaee (DL>.SO*I 1989) H^JEJ/ and F^GLSC^ (1976) consider that the finely reticulate grams present mm Cunomaceae [Vesseiowskya excepted) are unspecialized whereas grams with complete tectum and supratectaii processes are specialized conditions present in the grains of Saxifragaceae sensu stncto Thee present paper provides detailed information on She pollen grams of 24 species of Brunellia using both LMM and SEM Micrographs of pollen grams of some species of Cunomaceae Aistopetalum viticoides CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ &PAVÓN AND RELATED TAXA 33 3 3 £ 33 a: o: o: o: &; 33 3 1 o: Ï 3E att £L a: « ii-- a a_ u. S S tttt ir a 5 S LL 1 II I 66 « T5 5 (5 5 (MM <* >>. N*-3)(N*N*-fNifl l -- f^ #- ^ JI EN irt o o nn N n nn CM H nn IN ft CJ to o ^a ^a e** o* Ö DOOOOOOO O u u Tl l U. . li^ ^ 5 5 ££ c §§ 5. 44 1 CMM CNi CN 00 0 (VJ J en n( 0 0 0 0 rtrt ^ ^ T _^ « 88 5 i ÓÓ Ö Ó O O i/ï ÓÓO^OOotf>Ó(3C>*7T É óóóó6«iö'7 rjtöoaa o r^ ncN^P«(NT-rtioflr,T^'J1 " oT--flni-(D" aa Ï * , - ^^ 8. £ LftrvjS'ïii irt (O r ift ifl J: n ft* ro — oi if n r^ »-i3r ^* f^tDrNcn OO O ^ * *-_ r ^ O O - ^ q ^ Oi B ^ N W - r- crj O v- pï O O Cp ^- r-- liSSl l I'S ^'' ^ *- **' ^ 1P-" *-' T-' ,-" ^- ^ CT T-" O O" ^- T*" T-" 33 3 -£ aa OirtOO^»»nincDOOO»/>uï<3« ^ 11 f 1 óó ÖLftiTkOCióöóaiAo'óbóin ^rr 01 o — 11 * I O)) 3 S 4 cc S- * mm ^- 1- »n ^ ïiooïnoNnnooflSfflp)) in (oS'a'S'S'r^ ? sat t ^^ ^ n «- H 1-- T-N(N»-ï-»-rjl'J pp p m o * irr ^- g» 0) v aa (y- t öö *ti CN a> 3 oo 004rt*nirtOo *NN IN tn T- H MM hl ^ r ** * £ «« di in ó o -Is s dll 03 CD EÖ CHAPTERS CHAPTERS POLLENPOLLEN MORPHOLOGY OFBRUNELLIA RUIZ & PAVÖN AND RELATED TAXA 94 94 CaldcluvtaCaldcluvta pamculosa. Geissois superba, Lamanoma tomentosa were compared with grams of Bruneliia ass a close relationship was expected 6.. 2 MATERIALS AND METHODS Antherss from open flowers were removed from herbarium specimens ( COL. NY MO QCA and USi of 222 species of Bruneliia and of Spiraeanthemum katakata belonging to Cunomaceae (Table 6 1) Data of twoo additional species of Bruneliia previously studied were included in the present analysis (Oeczcc 1991)) Flowers of Acsnvthta were not available at that time Grains were acetolysed as outlined by Erdtmann ^1966) For SEM. pollen was mounted on stubs, coated with palladium gold and examined and photographedd with a Hitachi S-570 scanning electron microscope The species examined represent about halff of the total number of Bruneliia species and represent almost all of the clades from a phylogenetic analysiss of Bruneliia {Chapter 7). Slidess for LM were made with glycerine jelly and sealed with paraffin Sets of these preparations were depositedd in US and COL Measurements are based on at least 10 pollen grams (Table 6. 1 ) The size rangee and the mean values are given for each species Descriptions of pollen grains are based on observationss under LM and SEM. Shape classes are in accordance with ERDTMAN (1952) Pollen morphology terminologyy follows HIDEUX and FERGUSON (1976), PUNT et al. (1994) and NOWICKE (pers. comm ). 6.. 3 RESULTS 6.. 3.1 Pollen morphological descriptions BruneliiaBruneliia pollen grains are small to medium- sized (P X E = 11 0 - 32 5 x 10.5 - 28 0 um; Table 6. 1). Pollenn grain shape is prolate, subprolate, oblate spheroidal or occasionally suboblate (P/E = 0 88-1.48). Thee outline in polar view (amb) is (sub) circular. The grain is 3-colporate with 8.0- 25.0 um long colpi The endoaperturess are mostly elliptic pores, lalongate or lolongate {2.0-8 0 x 1 0-8.0 um ). The exine is incompletelyy (Fig 6 1a , b) to almost completely tectate (e. g. Fig 6 2). 6.. 3. 2 Categories of exine ornamentation Regardingg the exine ornamentation in Bruneliia. two extremes and three intermediate categories were distinguished.. One extreme is a reticulate tectum with large lumtna and high mun (Fig. 6 1 a, b). Due to thee interwoven aspect the term, reticulate striate could be applied to this ornamentation (Nowicke pers commm ). This type of ornamentation is the most incomplete (most open) and is observed in B comocladtfolia subsp.. comocladifolia Some species have a punctate tectum (Table 6.1) representing the other extreme, CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVON AND RELATED TAXA .-"" Fig.Fig. 6. 1 Scanning electron micrographs of Brunellia pollen . (a. b) B. comocladifolia ssp. comodadifolia. la) Equatorial view. (b) ExineExine ol striate reticulate type. (c. d) B. hygrothermica (c) Equatorial view, exine finely reticulate type, (d) Polar view showing circularcircular amb. (e, I) B comocladilolia ssp. cundinamarcencis (e). Equatorial view (!) Exine finely reticulate type, (g.h) B. dahenensis.dahenensis. (g) Ecuatorial view, (f) Finely reticulate type Scale bars 1pm CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVÓN AND RELATED TAXA 96 96 ass the tectum is almost complete (Fig. 6 2} The lumina are smaller and circular to slit-shaped This can bee observed in B susaconensis (Fig, 6. 2 a. b. OROZCO in prep.), S. brunnea (Fig. 6. 2 c. d) and S. costancensiscostancensis (Fig. 6. 2 e, f). Threee groups of species show intermediate forms between the two extremes of ornamentation. One intermediatee type is finely reticulate (Nowicke pers. comm.; Fig. 6. 1 c-h, Table 6. 1). The lumina can be recognizedd andthemun are joined at different levels, such as in S. comocladifolia subsp. cundinamarcensis (Figg 6. 1 e. f ). 6. dartenensis (Fig 6. 1 g, h). S hygrothermtca (Fig 6 1c d ). Another intermediate groupp includes species with muri and lumina of variable sizes with both muri and lumina at various levels (Figg 6 3 a-d) This type of sculpture is denoted as modified reticulate (Nowicke pers comm i The columellaee are visible as in S. glabra and B. goudotn (Fig 6. 3 a b) Columellae are not visible and the murii appear unevenly connected as in B. propinqua (Fig. 6. 3 c, d; Table 6 1} In a third group of species thee lumina are not distinct, and the muri are expanded, irregular m shape, and are connected at different levelss (Figs 6. 3 e, f: 6. 4 ) This sculpture type, modified rugulate ( Nowicke pers. comm ), is observed inn S boliviana, and in some populations of S. sibundoya (Fig. 6 3 f, Table 6. 1). A similar ornamentation typee is present in B inermis (Fig. 6. 4 a, b). B. cayambensis {Fig 6 4 c - e) and B weberbauen (Fig. 6.. 4 f). Ann overview of the ornamentation types of the exine in Bruneliia is presented in Figure 6 5 6.4.. DISCUSSION Pollenn morphological characters in Bruneliia indicate that despite variations of the exine ornamentation nonee of the grams is echinate or intectate. as was reported by BASAK (1967) and M^RT'CORE-JA (1970). respectivelyy Differences in the exine ornamentation of the pollen grains of Bruneliia were observed SEMM results of different populations of Bruneliia antioquensis. B comocladifolia and B sibundoya (OROZCOO 1991) partially covered the variation in exine ornamentation observed in the present study Two additionall groups of variation of the exine to those observed in OROZCO (1991) were seen the punctate typee with small, and circularto slit-shaped lumina (Fig 6 2) and the modified reticulate type (Fig 6 3a-d) Thee variations shown in OROZCO (1991). regular, irregular reticulate and rugulate perforate type correspond respectivelyy to Nowicke's terms striate reticulate {Fig. 6 1 a, b) finely reticulate (Fig 6 1 c, h). and rugulatee ornamentation (Figs 6 3 e, f; 6 4) Variations m the exine ornamentation were seen in populations off 6 sibundoya and B comocladifolia (OROZCO 1991) Finely reticulate (Fig. 6 1 c h) and rugulate CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVÓN AND RELATED TAXA Fig.Fig. 6. 2. Scanning electron micrographs of Brunelha pollen (a. b). B. susaconensis (a) Equatorial view, (b) Exine punctate type.type. (c. d). B. brunnea (c) Equatorial view, (d) Exme punctate type. (e. I) B. costaricensis. le) Equatorial view (f) Exme punctatepunctate type Scale bars 1fjm CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELLlA RUIZ &PAVÓN AND RELATED TAXA 98 98 ornamentationn (Figs. 6. 3 e, f; 6 4) were observed in different populations of the widely distributed species off S. sibundoya. (OROZCO. 1991. Fig. 10. p 982 versus Fig 11 p 9831 In different populations of S comocladifolia,comocladifolia, striate reticulate (Fig. 6. 1 a. b) and finely reticulate exme (Fig 6.1 e. f; were observed i ORCZZO,, 1991, Fig. 8, p 980 versus Fig 9. p.981) According to SARMIENTO (1986) the humidity related to thee altitude and temperature is an important factor in specific variation in the Colombian Andes. Thus, in thee case of intraspeciflc variation in the ornamentation of the exme and in accordance with the distribution off BruneHia in humrd montane forests, the humidity might be an influential factor. MULLER (1979) suggests thatt changes of exme sculpture m the same species are probably responses of the pollen gram to maintain aa stable pollen function Polymorphism in the ornamentation of the exme has been also found in other groupss as m Sapmdaceae. Dtmocarpus longan and species of Artyera i VAN DER HAMM 1993. VAN BERGEN F_-- 1995') They consider that in both cases the mtraspecific variation of the ornamentation is due to ecologicall causes and also related to the intraspecific variation of other characters. Thee number of BruneHia species studied under LM and SEM allows a characterization of variation in snapss and ornamentation of the pollen grains However relationships based on transmission electron Ticroscocicall data (TEM) are necessary for understanding the SEM results. The preliminary research uncerr ""EM analysis, applied to some species of BruneHia by NOWICKE {pers. cornm.) indicates that the variationn observed m SEM is due to variation in tectum thickness, and is not the result of longer columellae crr of a thicker foot layer The columellae m the grains of all taxa examined are uniformly short Using TEM analysis,, aiso a solid endexmous aperture membrane including a fineiy lamellate inner sublayer was ooserved.ooserved. TEM studies in BruneHia are necessary due to the polymorphism found in some species ndicatmgg that ornamentation is probably not a very conservative character for some species. Comcaredd to the pollen grains of BruneHia. Cunoniaceae pollen grains show much more variation, but aa reticulate exine pattern is usually present (HDEUX and FERGUSON 1976). Pollen grains are 2-3-colporate andd in some species as in Geissois superba (Fig. 6. 6 d ). 2-colpate grains are observed. In the available materiall of Cunoniacae 2-colporate grains are present in Lamanonia tomentosa (Fig. 6. 6 g) whereas. 3- colporatee grains occur in Aistopetalum viticoides, Caldcluvia paniculosa and Spiraeanthemun katakata (Fig.. 6 6 a. e, h). and 3-colporate grains are also present in Acsmithia pubescens. Dicolporate cr 2- colpatee pollen, presence of sexinous granules in some lumina of Geissois ternata (Fig.6 6 f), and the triangularr (amb) with apertures in the angles as in Aistopetalum viticoides, {Fig. 6. 6 h } are characters not presentt in BruneHia. CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVON AND RELATED TAXA ""-- % % % % Fig.Fig. 6 3. Scanning electron micrographs ofBrunellia pollen. (a) B. goudotn. equatorial view (b) B glabra, exme modified reticulate type.type. (c. d) B. propmqua. (c) Equatorial view, (d) Exme modified reticulate type, (e) B. boliviana equatorial view, (tj B sibundoyasibundoya ssp. sibundoya. exme rugulate type. Scale bars 1pm CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOGY OF BRUNELLlA RUIZ & PAVCN AND RELATED TAXA 100 100 NOWCKEE {pers comm ). with basis on a study of 12 species considers that the pollen of Cunomaceae iss fundamentally similar in sculpture However, the presence of 3-coiporate grams m Aistopeialum Catdcluvia,Catdcluvia, Sptraeantemum, and Acsmtthia and the generalized 2-colporate grains present in the remaining generaa (except Pullea glabra) might be related to the lack of monophyly in the family as was mentioned by HICKEYY and TAYLOR ( 1991) and also by OROZCO {1997) On the contrary, such variation might indicate polymorphismm as could occur in other characters The genera Aistopetalum, Spiraeanthemum and AcsmtthiaAcsmtthia closely related to Brunellia. (HUFFORD and DICKISON 1992) present 3-colporate grains Further researchh is necessary to establish whether the pollen morphology could be used as an informative character inn relationships within Cunomaceae. BRADFORD (pers. comm ) considered Cunomaceae as a monophyletic group,, based on molecular data. Displayingg the two states of pollen ornamentation used in the phylogenetic study of Brunellia (Chapter 7)) in the five categories, and superposing them on the cladogram it can be seen that the punctate exme withh circular to slit-like lumina (almost complete tectum) could be a piesiomorphic condition. This state is presentt in some species of Brunellia and Spiraeanthemun. with which Brunellia shares other characters (( DICKISON 1971,1980: EHRENDORFER ETAL 1984, CJATRECASAS 1985) The other exme categories incomplete tectumm (striate reticulate, modified reticulate and modified rugulate) are derived conditions The same couldd hold in Cunomaceae and Connaraceae. A complex exine sculpture, such as a rugulate ornamentation, presentt in Cunoniaceae (Vesselowskya and Pullea: HIDEUX and FERGUSON 1976) and also in Connaraceae (DICK.SONN 1980). is considered a specialized character In Brunellia the modified reticulate exme could be consideredd a synapomorphy of the Subsection Stmplicifoiiae (Chapter 7) m contrast to the homoplasic conditionn of the remaining categories of exme ornamentation in Brunellia. This modified reticulate condition iss very often related to foliar and inflorescence reduction and the presence of a navicular shaped endocarp (Chapterr 7). 6.. 5 ACKNOWLEDGEMENTS II thank RAYMOND VAN DER HAM from the Leiden branch of the National Herbarium of the Netherlands for hiss critical revision and suggestions. Thanks are also due to JOAN NOWICKE from the Smithsonian Institution forr guidance m the elaboration of LM and SEM microscope preparations and for providing some SEM micrographss of Cunomaceae pollen grains as well as for her critical revision Thanks also to my advisors HENRYY HGOGHIEMSTRA. from Amsterdam University, who encouraged me in the writing of this paper and PAUL JJ M MAAS from Utrecht University. This study was supported by the Smithsonian Institution in Washington DD C and the Instituto de Cienctas Naturales of the Universdad Nacional de Colombia and the Comité de Investigaciónn y Desarrollo Cientifico (CINDEC) of the Umversidad Nacional in Bogota for partially supporting CHAPTERS CHAPTERS POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVON AND RELATED TAXA 101 101 Fig.Fig. 6. 4. Scanning electron micrographs of Brunellia pollen . (a. b) B. inermis. la) Equatorial view, (b) Exine rugulate type, (c, d.d. e) B. cayambensis (c) Equatorial view, (d) Exine rugulate type, (e) Equatorial view, showing the endoaperture (!) B weberbaurei.weberbaurei. exine rugulate type. Scale bars 1pm. CHAPTERCHAPTER 6 POLLENPOLLEN MORPHOLOC Y OF BRUNELLIA RUIZ & PAVON AND RELATED TAXA 102 102 ** *\'?Sfa. n fa. '11 l F/g.. 6. 5. Summary of the exine ornamentation in Brunetlia. (a) B. comocladilolia ssp. comocladifolia striate reticulate exme. (b) B.B. comocladilolia ssp. cundinamarcensis, finely reticulate exine. (c) B. cayambensis. rugulate exine. (d) B. glabra, modified reticulatereticulate exine (e) B brunnea. punctate exine. Scale bars 1 urn. thiss research. I thank the Curators of all herbaria for providing the pollen material. I am especially grateful too SUSANN BRADEN from the Smithsonian Institution for her help in taking the SEM micrographs and to GIOVANII BOGOTA for the measurements of the pollen grains. I also thank GARY STILES from Institute de Cienciass Naturales for the revision of the original English version. Also thanks to the anonymous reviewers forr their critical comments. CHAPTER6 CHAPTER6 POLLENPOLLEN MORPHOLOGY OF BRUNELUA RUIZ& PAVON AND RELATED TAXA w'A w'A •• '^r d^hh v| I'^^K K^w^S rr 1 l^^»*fl lK-tiK J J M!^%H^ ^ ft ft Fig.. 6. 6 Scanning electron micrographs ol Cunoniaceae pollen, (a, b. c). Spiraeanthemum katakata. (a) Equatorial view, (b) Endoaperture.Endoaperture. (c) Exine punctate type, (d) Geissois superba. 2-colporate pollen, let Caidcluvia paniculosa. 3-colporate pollen. (!) GetssoisGetssois temata. granule in the exine punctate type (g) Lamanonia tomentosa. 2-colporate pollen (h) Aistopetalum viticoides. angu/aperturateangu/aperturate pollen. Scale bars 1 um. '~'~ 'CHAPTER 6 - _ - . P0LLEN MORPHOLOGY OF BRUNELUA RUIZ & PAVÓN AND RELATED'TAXA 104 104 SPECIMENSS INVESTIGATED BB boiivtana \Rusby) Cuatrec. BOLIVIA. La Paz. Nor-Yungas, La Paz 72 km toward Coroico, 28.8 1979. Beckk 1832. US 2232 COL. US (LM SMI SS brunnea Macbnde. PERU: Cuzco. Prov Paucartambo between Tambo of Tres Cruces andTambomayo. 144 5 Weberbauer 6969 US 2301 COL. US 1.30.1984.. Orozco &Tobón 1320 COL (LM, SM). S.. sibundoya Cuatrec. ssp sibundoya. COLOMBIA Cundinamarca Bojaca road, towards La Mercea 211 4 1983 Orozco & Lozano 1092 COL (LM. SM) SS stuebelu Cuatrec. COLOMBIA Cauca El Tambo Munchique 24 9 1983 Orozco et ai 1176 COL 2234COLUSS (LM. SM) 5.. subsesillis Killip & Cuatrec. COLOMBIA. Medellin. road to San Cristobal Cerro del Padre Amaya.11 11 1985. Orozco et al 1425 COL 2302 COL. US (LM.SM) 8.8. susaconensis (Cuatrec.) Orozco. COLOMBIA: Boyaca. Susacbn. Santa Rosita to Onzaga road.. 10 1987. Orozco et al 1810 COL 2296 COL, US (LM.SM) 88 tomentosaH &B ECUADOR Pichincha, below San Juan towards Chtnboga 2 8. 1955 Asplundd 17162 NY 2231 COL, US (LM. SM) B.. trianae Cuatrec. COLOMBIA: Antioqura, Medellin. 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