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Pollen Ultrastructure of the Biovulate Euphorbiaceae Author(S): Michael G

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Pollen Ultrastructure of the Biovulate Euphorbiaceae Author(S): Michael G Pollen Ultrastructure of the Biovulate Euphorbiaceae Author(s): Michael G. Simpson and Geoffrey A. Levin Reviewed work(s): Source: International Journal of Plant Sciences, Vol. 155, No. 3 (May, 1994), pp. 313-341 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2475184 . Accessed: 26/07/2012 14:35 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 [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. http://www.jstor.org Int.J. Plant Sci. 155(3):313-341.1994. ? 1994by The Universityof Chicago. All rightsreserved. 1058-5893/94/5503-0008$02.00 POLLENULTRASTRUCTURE OF THE BIOVULATE EUPHORBIACEAE MICHAEL G. SIMPSON AND GEOFFREY A. LEVIN' Departmentof Biology,San Diego StateUniversity, San Diego,California 92182-0057; and BotanyDepartment, San Diego NaturalHistory Museum, P.O. Box 1390,San Diego,California 92112 Pollenultrastructure of the biovulate Euphorbiaceae, including the subfamilies Phyllanthoideae and Oldfieldioideae,was investigatedwith light, scanning electron, and transmissionelectron microscopy. Pollenof Phyllanthoideae, represented by 12 speciesin ninegenera, was prolateto oblate,almost always 3-colporate,rarely 3-porate or pantoporate,and mostlywith reticulate, rarely baculate, echinate, or scabrate,sculpturing. The ektexineshowed little variation in ultrastructure;almost without exception it was tectate-columellatewith a homogeneousfoot layer, columellae continuous with both the tectum and thefoot layer, and a perforate,homogeneous tectum. The onlymajor exception was Amanoa guianensis, whichwas intectateand baculateas theresult of the evolutionary loss of thetectum. Pollen of Oldfiel- dioideae,in whichwe studied28 speciesfrom 25 genera,and obtaineddata fromthe literature for two of thethree remaining genera in the subfamily,was globoseto oblate,with four or more,rarely only three,apertures that were either equatorial and brevicolporate toporate or pantoporate, and with echinate, rarelyscabrate, sculpturing. The ektexineshowed considerable variation in ultrastructure.The most widespreadstructural type consisted of a footlayer that is eitherthin and irregular or absent, an interstitium composedof irregular to columellateexine deposits that are discontinuous from the foot layer, if present, and a tectumcomposed of laterallyappressed baculate elements that delimit fine microperforations. A secondcommon structural type differed in havinga relativelythick, homogeneous foot layer and thin, granularto tabular,microperforate tectum. Isolated genera have a reducedinterstitium and/or a homo- geneoustectum or show a reversalto a tectate-columellatearchitecture. Variation in aperture morphology, sculpturing,and exineultrastructure is phylogenetically significant in theOldfieldioideae. Introduction remainingthree subfamilies share the synapo- Palynologicalstudies of the Euphorbiaceae have morphy(shared derived character state) of a sin- beenextremely useful in clarifyingsystematic re- gleovule per locule. The Acalyphoideae (113 gen- lationshipswithin the family.Using lightmi- era, 2,100 species) is probablyparaphyletic, croscopy(LM), Erdtman(1952) was the firstto differingfrom the otheruniovulate subfamilies pointout the great variation in theshape, sculp- mainlyin lackingmilky or coloredlatex. The turing,and apertures of Euphorbiaceae pollen and Crotonoideae(64 genera,1,450 species) share the to suggestthat these characteristics were signifi- synapomorphiesof stellate, dendritic, or lepidote cant systematically.Punt (1962) subsequently trichomes,and mostlyporate or inaperturate pol- conducteda more detailedsurvey, confirming len withthe distinctive"crotonoid" pattern of most of Erdtman'sconclusions. K6hler (1965) exinousprocesses (Nowicke 1994). The Euphor- completeda thoroughstudy of the biovulate gen- bioideae(38 genera,1,800 species) are unitedby era,i.e., those that produce two ovules per locule, thesynapomorphy of a reducedperianth. whichhad long been placed in the subfamily Our understandingof relationships within the Phyllanthoideae.He concludedthat the genera Phyllanthoideaeand Oldfieldioideaehas been withspinose pollen shouldbe placed in a new greatlyincreased by pollen studiesusing SEM subfamily,the Oldfieldioideae. (Bonnefilleand Riollet 1980; Punt 1980, 1987; Subsequentintrafamilial classifications of the Poole 1981; Haydenet al. 1984;Webster 1984a; Euphorbiaceaehave continued to recognizethese Dechampset al. 1985; El-Ghazalyand Raj 1986; two subfamiliesas well as the Acalyphoideae, Lobreau-Callenin McPhersonand Tirel 1987; Crotonoideae,and Euphorbioideae(Webster Websteret al. 1987;Lobreau-Callen and Suarez- 1967,1975, 1994).The Phyllanthoideae(55 gen- Cervera1989). However,only three palynologi- era,2,100 species)is probablya basal, paraphy- cal studies of these subfamilies(Poole 1981; letictaxon, the members of which are united only Haydenet al. 1984; Lobreau-Callenand Suarez- bysharing a numberof ancestral (plesiomorphic) Cervera1989) have used TEM to describethe features.The Oldfieldioideae(28 genera,80 spe- detailsof exinearchitecture, and thoughthe au- cies) appearsto be monophyletic,united by the thorstreated only six genera,they suggested that spinosepollen observed by Kohler(1965). The pollenwall ultrastructuremay be systematically valuable. We investigatedpollen ultrastructurein the ' Authorfor correspondence and reprints. Current address: biovulateEuphorbiaceae with the goal of ex- Centerfor Biodiversity, Illinois Natural History Survey, 607 ploringits contribution to thesystematics of these East PeabodyDrive, Champaign, Illinois 61820. subfamilies.We initiallyfound little variation in Manuscriptreceived September 1993; revisedmanuscript re- exinewall structurewithin the Phyllanthoideae, ceivedJanuary 1994. whereaspollen of Oldfieldioideae varied consid- 313 314 INTERNATIONAL JOURNAL OF PLANT SCIENCES erably.As a consequencewe concentratedour ips EM 410 transmissionelectron microscope at effortson thelatter subfamily, examining a small 80 kV. sampleof Phyllanthoideaeselected to included Mean pollengrain diameter (polar x equato- membersof the major groups recognizedin rial)was determined from LM preparations,mean Webster'sclassification (1975, 1994).We also ex- spinelength from either LM observationsor TEM aminedDidymocistus and Hymenocardia,both micrographs,and mean ektexinefoot layer and Phyllanthoideae,because of questions about their tectumthicknesses from TEM micrographs. relationships. Samplesizes fromLM observationsincluded at This articleis primarilydescriptive. In a sep- least 10 pollengrains, whereas those from TEM aratearticle (Levin and Simpson1994a), we an- micrographswere generally very small. alyzethe pollen characters of the subfamily Old- Our terminologygenerally follows that of fieldioideaephylogenetically, both alone and in Reitsma(1970) and Walkerand Doyle (1975). combinationwith anatomical and morphological Reitsma(1970) distinguishedspines of length > 1 characters.Only the major conclusionsof our ,um,which he termed"echinae" (sculpturing phylogeneticanalysis will be repeatedhere. "echinate"),from those of length < 1 ,um,which he termed"scabrae" (sculpturing "scabrate"). To reflectthe discontinuities among the taxa in our Materialand methods study,we modifiedReitsma's terms and use the Flowersfrom dried herbarium specimens were terms"echinae" or "echinate"when spines are rehydratedeither at roomtemperature or at 600C >0.9 ,umin lengthand "scabrae" or "scabrate" in 1%Aerosol OT for3-5 d,then fixed and stored when <0.5 ,umin length.With regard to tectal in FAA. For LM, pollenwas clearedin Hoyer's structure,we use the term"microperforate" to mountingmedium (Radford et al. 1974) on a describea tectumpenetrated by very narrow pores microscopeslide preparation.LM observations delimitedby closelyappressed tectal elements, weremade with differential interference contrast whereas"homogeneous" refers to theabsence of opticsusing a Nikon Microphot-FXphotomi- microperforationsor any otherwall discontinu- croscope.For SEM, whole antherswere dehy- ities.A microperforatetectum may be composed dratedto 100%ethanol, then gradually infiltrated eitherof laterally appressed baculate elements, a to 100% Freon 113 or methylal(dimethoxyme- structurethat we term "microperforate/bacu- thane).The antherswere placed in a metalcap- late,"or oflaterally appressed granular to tabular suleand critical-pointdried with a Tousimiscrit- elements,a structurethat we term"microperfor- ical-pointdryer using pressurized carbon dioxide ate/granular.""Perforate" refers to a largerhole as thetransition fluid. Pollen grains were tapped in the tectumthat is not immediatelybordered ontoa stubcovered with double stick tape, sput- by closelyappressed tectal elements; the tectum ter-coatedwith gold/palladium in a Hummer-4 betweenperforations may be eithermicroperfor- sputteringapparatus, and photographedon a Hi- ateor homogeneous. "Imperforate" simply means tachiS500 scanningelectron microscope (20 kV). thatthe tectum lacks both perforations and mi- In additionto
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