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Septal Nectary Anatomy and Phylogeny of the Haemodoraceae Author(S): Michael G

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Septal Nectary Anatomy and Phylogeny of the Haemodoraceae Author(S): Michael G Septal Nectary Anatomy and Phylogeny of the Haemodoraceae Author(s): Michael G. Simpson Source: Systematic Botany, Vol. 18, No. 4 (Oct. - Dec., 1993), pp. 593-613 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/2419536 . Accessed: 13/04/2011 15:47 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=aspt. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. 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]. American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany. http://www.jstor.org SystematicBotany (1993), 18(4): pp. 593-613 ? Copyright 1993 by the American Society of Plant Taxonomists Septal Nectary Anatomy and Phylogeny of the Haemodoraceae MICHAEL G. SIMPSON Department of Biology, San Diego State University, San Diego, California 92182-0057 ABSTRACT. Septal nectaryanatomy of membersof the Haemodoraceae is described,with emphasis on nectarynumber and relativeposition within the ovary.Three typesof septal nectariesare defined: infralocular,interlocular, and supralocular. The phylogenetic and possible adaptive significanceof these featuresare assessed by adding the data to a previous cladistic analysis of the Haemodoraceae and considering cladistic patternsin termsof functionalfloral morphology. I hypothesize that: 1) three interlocular septal nectaries are ancestral for the Haemodoraceae, but were secondarily ac- quired in the genus Anigozanthos(Conostylideae) in response to selective pressure for increased nectar production forbird pollination; 2) three supralocular nectariesconstitute a synapomorphy for all or most of the tribe Conostylideae, but evolved independently in the genus Dilatrisof the Haemodoreae; 3) two infralocularnectaries evolved concomitantlywith unique "perianth aper- tures" and arose via the evolution of zygomorphy and basal displacement of nectaries into the receptaculartissue; and 4) septal nectaries were independently lost in the genera Xiphidiumand Phlebocarya,perhaps in response to a shiftin pollination mechanism. The Hae-modoraceae are a monophyletic I recently analyzed the phylogenetic rela- group consisting of 13 currently recognized tionshipsof genera and species complexes with- genera and approximately100 species (Ander- in the Haemodoraceae and proposed some berg and Eldenas 1991; MacFarlane et al. 1987; changes in classification(Simpson 1990). In or- Simpson 1990); the monotypicMacropidia fuligi- der to understand better the evolution of the nosa has recentlybeen reclassifiedas a member diverse floralmorphology in the group and to of the genus Anigozanthosby Anderberg and help resolve incompatibilities of certain hy- Eldenas (1991), a viewpoint with which I whol- potheses of characterevolution, I have initiated ly agree. Members of the Haemodoraceae are studies of floral anatomy, ovary development, rhizomatous to stoloniferous herbs, with eq- and ovule/seed developmentin the family.This uitant,unifacial leaves and mostly cymose in- paper describes observations on the anatomy florescences and have distributionsspanning and spatial relationship of septal nectaries in Australia, South Africa, South and Central the Haemodoraceae (and near outgroups) and America,and eastern North America. The fam- the phylogeneticsignificance of these new data. ily exhibitsconsiderable diversityin floralmor- More detailed anatomical descriptionsof septal phology. Flowers range in size from approxi- nectaryepithelial cells, subepithelial cells, and mately5 mm long in the bee-pollinated genera surrounding vasculature and ergastic sub- Xiphidium(South and Central America) and stances will appear in another publication. Phlebocarya(southeastern Australia) to over 6 Dahlgren and Clifford(1982) reviewed the cm long in species of the bird-pollinated Ani- occurrence of floral nectaries in monocotyle- gozanthos(southwestern Australia). Flowers may dons, relying primarilyon the work by Dau- be either actinomorphic or zygomorphic, en- mann (1970). The three major types of nectaries antiostylous or symmetrically-styled,glabrous in monocots that they defined are: 1) perigo- or densely tomentose (ranging in color from nal-nectaries at the adaxial bases of tepals; 2) white, yellow, green, orange, red, to black); the androecial-nectaries on filamentsof stamens perianthcan be either imbricateor valvate; sta- or staminodes; and 3) septal-nectaries found men number is 6, 3, or 1; ovary position is su- in the septal regions of the gynoecium. Rare perior,half-inferior, or inferior;locule number typesof monocot nectariesinclude those found and fertilecarpel number is either 3 or 1; and at stylebases, stigmas,surface of carpels or ova- ovules and seeds may be 1, 2, 4, 5-7, or indef- ry,and nectariferousdisks. Septal nectariesare inite in number,epitropous or hypotropousin by far the most common type found in mono- position,and considerablyvariable in shape and cotyledons and are entirely lacking in dicots size (see Simpson 1990). (Schmid 1985). Septal nectaries are nectar-se- 593 594 SYSTEMATIC BOTANY [Volume 18 cretingcavities that occur in the septal region Commeliniflorae,sensu Dahlgren and Clifford between adjacent carpels by the incomplete fu- 1982), septal nectaries are absent. Only Com- sion of carpels during the development of the melinais reportedto have nectariesat all, these gynoecium ("lack of intercarpellarypostgenital being ofthe androecial type(Dahlgren and Clif- fusion,"Schmid 1985). Smetsand Cresens (1988) ford1982). Among otherputative close relatives proposed that the name "gynopleural" nectary of the Haemodoraceae, (sensu Clark et al. 1993; replace that of "septal" nectaryin order to in- Dahlgren and Clifford 1982), septal nectaries dicate possible homologybetween nectariesthat are widespread in the Bromeliaceae and Zin- occur on outer carpellarysurfaces and those oc- giberales. In the Bromeliaceae, septal nectaries curringinternally. However, I choose to retain have been reported in 35 investigated genera the term"septal" nectaryin the present paper (ca. 90 species) of all three subfamilies (Bohme because of its widespread use in the literature. 1988). Among these taxa, septal nectaryanato- The cavities of septal nectaries are generally my is quite diverse; the position of the nectaries verticallyoriented and are delimited by one or is eitherbelow (equivalent to "infralocular"used more layers of nectar-secretingepithelial cells here), within (equivalent to "interlocular"used and subepithelial cells. The nectaries may be here), or above (equivalent to "supralocular" foundin eithersuperior- or inferior-ovariedtaxa used here) the locular region of the ovary. In and secrete nectar via pores or slits at the base the Zingiberales, all members of which have or apex of the ovary (Daumann 1970). Schmid predominantlyinferior ovaries, septal nectaries (1985) recognized morphological criteria for are known to occur in seven of the eight fam- distinguishingseptal nectarytypes, among these ilies (lacking only in Lowiaceae). Of those fam- being: transectionaloutline (including whether ilies possessing septal nectaries, nectary posi- simple, or labyrinthine/convoluted),distinc- tion varies. The nectaries occur within the tiveness (whether unfused or confluent),ver- locular region (equivalent to "interlocular"used tical extent,distance from ovarian center,and here) in members of the Cannaceae, Heliconi- position of opening in gynoecium. aceae, Marantaceae, and Strelitziaceae (a por- In the Haemodoraceae, septal nectaries have tion of the nectaries also occurring below the been reported in all but one of the 13 genera locules in the Heliconiaceae); nectaries occur defined here (Dahlgren and Clifford 1982; above the locules (equivalent to "supralocular" Simpson 1990). The closest relativesto the Hae- used here) in members of the Costaceae, Mu- modoraceae have recentlybeen considered to saceae, and Zingiberaceae (Kirchoff, pers. be the Pontederiaceae and Typhales, with the comm.; Newman and Kirchoff1992). Although Philydraceae, Bromeliaceae, Velloziaceae, and the Bromeliaceae and Zingiberales were not Zingiberales, respectively, more distantly re- designated as outgroupsin the presentcladistic lated (Dahlgren and Rasmussen 1983). [The Vel- study, they should be considered in broader loziaceae have been suggested to be ratherdis- phylogenetic analyses of the complex. tant relatives to the Haemodoraceae based on chloroplast DNA rbcL sequence data (Clark et MATERIALS AND METHODS al. 1993).] Simpson (1987, 1990) argued that the Pontederiaceae are the most likely sistergroup Live, maturebuds
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