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Morphological Variation of Some Floral Features of the Subfamily

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Morphological Variation of Some Floral Features of the Subfamily BOT.GAZ. 149(1):82-91. 1988. t 1988 by The Universityof Chicago. All rightsreserved. 0006-807 1/88/490 1-0008$02 .00 MORPHOLOGICALVARIATION OF SOMEFLORAL FEATURES OF THE SUBFAMILY PITCAIRNIOIDEAE(BROMELIACEAE) AND THEIRSIGNIFICANCE IN POLLINATIONBIOLOGY G. S. VARADARAJANAND G. K. BROWN Departmentof Botanyand MarionOwnbey Herbarium, Washington State University,Pullman, Washington 99164-4230; and Departmentof Botany, Universityof Wyoming,Laramie, Wyoming 82071 Scanningelectron and light microscopyobservations of wet-preservedflowers of Bromeliaceaesubfamily Pitcairnioideaeyield new informationon the stigma, petal scales, and septal nectaries.Variations of the stigmatypes are evidentamong several genera. The gross structuralfeatures of the stigmado not indicate definitepollination trends, but the shape of the lobes and papillaeindicate a few specific modes. In pit- cairnioidgenera, petal scales, when functional,may aid in pollinationby accumulatingthe nectarsecreted from the ovary, thus facilitatingits availabilityto the pollinator.Nectaries associated with the gynoecia usuallydisplay tripartite channels in the ovarysepta. Some developmentalchanges of the channelstructure andposition of the ovaryindicate three probable modes of nectarrelease from the gynoeciaof the pitcair- nioids:(1) throughlateral grooves or openings,(2) partlythrough the apicalorifices and partlythrough the dissolvedareas of the spetal channels,and (3) exclusively throughthe apical orifices. Analysis of a wide rangeof floral featuresindicates that ornithophily, chiropterophily, and entomophilyexist in differentPit- cairnioideaelineages. Introduction Gynoecia of the monocotyledons often contain nectaries in the ovary septa (septal nectaries) and lnvestigations on Bromeliaceae have produced are known to be of taxonomic value in several fam- descriptive and systematically useful data (BROWN ilies (DAUMANN1970; FAHN1982; THORNE1983; and GILMARTIN 1984;BROWN et al. 1984;VARA DAHLGRENet al. 1985) . BUDNOWSKIS ( 1922) ex- DARAJANand BROWN 1985;GILMARTIN and BROWN amination represents the only major study of Bro- 1985,1986;VARADARAJAN 1986a; VARADARAJAN meliaceae septal nectaries. Their morphological and GILMARTIN 1988),and because of its primitive variation and the significance in pollination were status, the subfamily Pitcairnioideae was a major not analyzed until the present work. focus (SMITH 1934;MARCHANT 1967;TOMLINSON The goal of this study is to examine the diversity 1969;BENZING 1980;GILMARTIN and BROVVN1987). of stigma morphology, petal scales, and septal nec- Mostly, the gynoecial traits of Bromeliaceae were taries of the subfamily Pitcairnioideae from wet- described, with a little information on the floral preserved material. features of the Pitcairnioideae (GARDNER 1982; UTLEY 1983;BROWN and GILMARTIN 1984;GIL Material and methods MARTIN and BROWN 1985). The 41 species of nine genera of Pitcairnioideae Several floral characteristics (petal scales, nec- were examinedby light microscopy;30 species were taries) were described only from dried and rehy- also examined with scanning electron microscopy drated material, which vwas partly responsible for (SEM) (table 1). All material was field-collected the poor structural detail of floral traits. Petal scales by G. S. VARADARAJANand preserved in FAA (ligule, petal appendages, nectariferous scales, lat- (formaldehyde:acetic acid:ethanol, 1: 1: 18). Sam- eral folds, vertical calli) are relatively frequent in ples of stigmas and ovaries were selected from pre- Bromeliaceae and are diagnostic of several genera anthesis to post-anthesis, and the internal structure (MEZ 1934- 35; SMITH and DOWNS 1974, 1977, of ovaries was examined from freehand cross sec- 1979).Aside from the taxonomic value associated tions. Petal scales were examined from nearly ma- with petal scales, their morphological variation and ture (anthesis) flowers. significance in pollination were not well eluci- Floral parts selected for SEM were transferred dated. from FAA to 0.1 M cacodylate buffer, postfixed in OSO4, dehydrated through an ethanolic series (30%, 50%, 70%, 95%, 100%), dried in a Bomar Manuscript received September 1986; revised manuscript re- critical-point dryer, mounted on aluminum stubs, ceived February 1987. and gold coated with a Technics Hummer-Sputter Address for correspondence and reprints: G. S. VARADARA coater. Flower samples were then examined in an JAN,Arnold Arboretum, Harvard University, 22 Divinity Ave- ETEC Auto Scan U- 1 scanning electron micro- nue, Cambridge, Massachusetts 02138. scope at 20 kV. 82 VARADARAJAN& BROWN FLORALFEATURES OF PITCAIRNIOIDEAE 83 Results and discussion in Brocchinia,Cottendorfa, and Fosterella. The convolute-bladestigma, type III, previously re- FLORALFEATURES OF PITCAIRNIOIDEAE ported only from the subfamily Tillandsioideae STIGMAMORPHOLOGY. Terminology of the occursin Navia (fig. 7). stigma types follows BROUNand GILMARTIN(1984). Stigmatypes vary within themselvesas well as The most commonly observed stigma type in the withincertain taxa; e.g., species of Brocchiniaex- subfamily is the conduplicate-spiralpattern, type II hibit types I and II (table 2). We confirmreports (f1gS. 1-6, 8-11). The simple-erect type I occurs of a modifiedconduplicate-spiral type (type II) in ,& J 7g:> t- w i58 w * L'< '\C'4-'X,w. t-. FIGS. 1-5.-SEM of variationin stigmamorphology. Fig. 1, Ayensuauaipanensis (Varadarajan 1196). Type II stigma. Arrow indicatesasymmetric lobe of stigma.Note bulbouspapillae (BP) and spiralingrestricted to individuallobes. Figs. 2, 3, Brocchinia acuminata(Varadarajan & Oliva 1159). Two views of a type II stigma. Note tubular,branched papillae (TP). Fig. 4, Dyckia brevifolia(SEL 82-558). Type II stigma at pre-anthesis.Arrows indicate highly compactstigma lobes. Fig. 5, Deuterocohnia schreiteri(Varadarajan et al. 1247). Type II stigmaduring post-anthesis. Note pollen grainstrapped within dissected lobes (DL) of stigmabranches. Compare with papillaein othertaxa. r* %l, t ( , s x - . .S. *: w @w t l t :.# .: |oE L.d i s: f t.: 'R....f:nl::N 4 ,La. r; ,. l. .| *::.,\|' :.sl ::l :::|::||:l . A-==at SslisK Type II stigma at post- Dyckia ragonesei (Varadarajan1218). in stigma morphology.Fig. 6, pollen (P). Fig. 7, Navia splen- FIGS. 6-9. SEM of variation (DL) of stigmalobes with stigmaticlobes. Note dissectedareas Pitcairnia nuda (SEL 81-1979). anthesis,displaying the loosened convoluteblades (CB); Fig. 8, 9, Type III stigma duringanthesis with papillateoutgrowths (TP). Fig. dens (Varadarajan1215). mostly the individuallobes. Tubular lobes and duringanthesis. Spiral folding involves noticeablespiral folding of individual Type II stigma Type II stigmaduring anthesis with heterophylla(Varadarajan 1171). of stigma lobes. Pitcairnia papillae(BP) are confinedto edges entirestigmatic apparatus. Bulbous ,wr};9, ;fX 0 ;. 0 0 8.;.:f f X 20'0,llmX < EEWS >E;f2X f# rl^2"tWt VARADARAJAN& BROWN-FLORAL FEATURESOF PlTCAIRNIOIDEAE 85 FIGS.10-13. Figs. 10, 11, SEM of variation in stigma morphology in Puya. Fig. 10, P. Iaxa (SEL 83-207). A pre-anthesis type II stigma. Arrow indicates stigma lobes that begin to undergo spiraling. Papillae absent at this stage. Fig. 11, P. harmsii (Varadarajanet al. 1245). Type II stigma with highly compacted, spirally folded lobes at anthesis. Bulbous papillae (BP) are confined to edges of stigma lobes. Figs. 12, 13, SEM of petal scales of Deuterocohnia.Note the similarity of scales within the two species. Fig. 12, D. Iongipetala(Varadarajan et al. 1244). Fig. 13, D. haumanEi(Varadarajan et al. 1250). B. acuminata(figs. 2, 3) and of a weakly condu- of twistedor spiraledbranches, (3) durationof the plicate-spiral type in B. gilmartinii(VARADARAJAN spiralfolding, and (4) shape and symmetryof the 1986b). The stigma of B. steyermarkiiis similar to individuallobes. type I reported in B. reductaBaker (BROWNand PAPILLAE.Stigma lobes of some taxa are as- GILMARTIN1984). sociated with relatively small, simple, bulbous Variability in types I and II is evident with re- structuresknown as the papillae (fig. 11). These gard to the (1) papillae on the lobes, (2) presence are occasionally tubular (fig. 8), complex, and N.splendens L.B. Smitha ................... TABLE 1 TAXA INCLUDEDIN THESTUDY OF FLORAL FEATURES OF THE SUBFAMILY PITCAIRNIOIDEAE Taxa Details of voucher Locality Ayensua: A. uaipanensisL. B. Smitha..... Varadarajan1 196 Venezuela:Bolivar, AuyanTepui Brocchinia: B. acuminataL. B. Smitha...... Varadarajan& Oliva 1159 Venezuela:Bolivar, La Escalera B. gilmartinfiVaradarajana ...... Varadarajan& Oliva 1158 Venezuela:Bolivar, La Escalera B. steyermarkiiL. B. Smitha..... Varadarajan& Oliva 1164 Venezuela:Bolivar, La GranSabana Cottendorfia: C. guianensisK1. ex Bak.a ...... Varadarajan& Oliva 1161 Venezuela:Bolivar, Kavanayan Dezaterocohnia: D. haumaniiCastell.a........... Varadarajanet al. 1250 Argentina:Salta, Cafayate D. IongipetalaMeza ............ Varadarajanet al. 1244 Argentina:Cordoba, Sierra Chica D. schreiteriCastell.a........... Varadarajanet al. 1248 Argentina:Salta, Cafayate Dyckia: D. brevifoliaBak.a ............. SEL 82-558b Brazil D. niederleiniMeza ............ GSV 0030b Brazil D. ragoneseiCastell.a .......... Varadarajan1218 Argentina:Entre Rios Rio Parana Fosterella: F. elata Luthera............... Holotypeb Bolivia E. pendulifloraL. B. Smitha..... SEL 024397b Peru Navia. Varadarajan1215 Venezuela:Bolivar, AuyanTepui Pitcairnia: P. andreanaLindena ........... SEL 82-554b Colombia P. armataMaurya.............. Varadarajan& Gaunchez 1150 Venezuela:.T.F. Amazonas,
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