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Pollination Drop in Relation to Cone Morphology in Podocarpaceae: a Novel Reproductive Mechanism Author(S): P

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Pollination Drop in Relation to Cone Morphology in Podocarpaceae: a Novel Reproductive Mechanism Author(S): P Pollination Drop in Relation to Cone Morphology in Podocarpaceae: A Novel Reproductive Mechanism Author(s): P. B. Tomlinson, J. E. Braggins, J. A. Rattenbury Source: American Journal of Botany, Vol. 78, No. 9 (Sep., 1991), pp. 1289-1303 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/2444932 . Accessed: 23/08/2011 15:47 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]. Botanical Society of America is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany. http://www.jstor.org AmericanJournal of Botany 78(9): 1289-1303. 1991. POLLINATION DROP IN RELATION TO CONE MORPHOLOGY IN PODOCARPACEAE: A NOVEL REPRODUCTIVE MECHANISM' P. B. TOMLINSON,2'4 J. E. BRAGGINS,3 AND J. A. RATTENBURY3 2HarvardForest, Petersham, Massachusetts 01366; and 3Departmentof Botany, University of Auckland, Auckland, New Zealand Observationof ovulatecones at thetime of pollinationin the southernconiferous family Podocarpaceaedemonstrates a distinctivemethod of pollencapture, involving an extended pollinationdrop. Ovules in all generaof the family are orthotropousand singlewithin the axil of each fertilebract. In Microstrobusand Phyllocladusovules are-erect (i.e., the micropyle directedaway from the cone axis) and are notassociated with an ovule-supportingstructure (epimatium).Pollen in thesetwo genera must land directly on thepollination drop in theway usualfor gymnosperms, as observed in Phyllocladus.In all othergenera, the ovule is inverted (i.e., the micropyleis directedtoward the cone axis) and supportedby a specializedovule- supportingstructure (epimatium). In Saxegothaeathere is no pollinationdrop and gametesare deliveredto theovule by pollen tube growth. Pollination drops were observed in sevenof the remaininggenera. In thesegenera the drop extends over the adjacent bract surface or coneaxis and can retainpollen that has arrivedprior to dropsecretion ("pollen scavenging"). The pollen floatsupward into the micropylar cavity. The configurationofthe cone in othergenera in which a pollinationdrop has notyet been observed directly suggests that pollen scavenging is general withinthe family and mayincrease pollination efficiency by extendingpollination in spaceand time.Increased pollination efficiency may relate to thereduction of ovule number in eachcone, oftento one in manygenera, a derivedcondition. A biologicalperspective suggests that animal dispersalof large seeds may be theultimate adaptive driving force that has generatedthe need forgreater pollination efficiency. We heredocument that the pollination drop fleshyseeds dispersedby animals (mainly mechanismof many members of the southern birds). hemisphereconiferous family Podocarpaceae A pollinationdrop is knownto be produced (podocarps)differs from that in most gym- in all speciesof Cephalotaxaceae,Cupressa- nosperms.The dropcan "scavenge"pollen that ceae,Taxaceae, and Taxodiaceae that have been has previouslyfallen near the ovule,prior to studied(Takaso, 1990). In Araucariaceaeno dropsecretion. Presumably this increases the pollinationdrop occurs, the gametes being de- efficiencyof pollinationand is correlatedwith liveredto themicropyle by pollen tube growth thereduced number of ovules per cone in most (Eames, 1913; Haines, Prakash,and Nikles, podocarps.The Podocarpaceae,unlike most 1984). The greatestdiversity in pollination otherconifers, are largelycharacterized by mechanismsis knownfor Pinaceae, in which a pollinationdrop is reportedfor Picea and Pinus,while othergenera have mechanisms 1 Receivedfor publication 21 January1991; revision withoutdrop secretion, such as pollenengulf- accepted7 May 1991. The authorsthank the Directors and ChiefRangers of mentor pollen tube growth (Doyle, 1945). Ab- Mt. TongariroNational Park; AucklandRegional Au- senceof a dropis consideredderived since the thorityBotanical Garden, Manurewa; Waitakere Conser- presenceof a pollinationdrop may be a very vationArea, Auckland; Kauaeranga Forest Reserve, Cor- ancientfeature of gymnosperms (cf. Rothwell, omandel,all in New Zealand;Mt. Field NationalPark, 1977). In Podocarpaceaethere has been no Tasmania;and Fairchild Tropical Garden, Miami, Florida in forpermission to collect and makeobservations in several observationof pollination mechanisms nat- protectedareas; Alison Shapcott, University of Tasmania; urallygrowing trees. Doyle (1945) suggested C. J.Quinn, University ofNew South Wales; J.-M. Veillon, thatthe Podocarpaceae formed a parallelseries CentreO.R.S.T.O.M., Noumea, New Caledonia; and Ewen to the Pinaceae,also involvingloss of a pol- K. Cameron,University of Aucklandfor further collec- linationdrop, but he describeda pollination tionsand fieldassistance; and EllenMoriarty and Toku- drop onlyin one specieseach of Podocarpus shiroTakaso for help with illustrations. Field work in the SouthPacific was supportedby a grantto PBT fromthe and Prumnopityscultivated in Irishgardens. ResearchCommittee of the National Geographic Society. Saxegothaea,which lacks a pollinationdrop 4Author forcorrespondence. (Noren,1908; Doyle and O'Leary,1935), was 1289 1290 AMERICAN JOURNAL OF BOTANY [Vol. 78 TABLE 1. Podocarpaceae.Taxa withan observedpollination drop No. fertile Taxon Localityr bracts Ovule orientationb Dacrycarpusdacrydioides Huia (NZ) 1(-2) Inverted Halocarpus kirkii Kelly's Road, Waitemata (NZ) 2-3 Inverted Lagarostroboscolensoi Cult. Auckland Univ. Campus (NZ) 2-3 Inverted Lepidothamnusintermedius Kauaeranga For. Res. (NZ) 1(-2) Obliquely erect Lepidothamnuslaxifolius TongariroNat. Park (NZ) 1(-2) Obliquely erect Microcachrystetragona Mt. Field Nat. Park (T) 8-16 Inverted Phyllocladusaspleniifolius var. alpinus TongariroNat. Park (NZ) 1-5 Erect Phyllocladustrichomanoides Cornwall Park, Auckland (NZ) 2-7 Erect Podocarpus macrophyllus Cult. Fairchild Tropical Garden (US) 1-2 Inverted Podocarpus nivalis TongariroNat. Park (NZ) 1-2 Inverted Podocarpus totara Manurewa Bot. Gard. (NZ) 1-2 Inverted Prumnopitysandina Cult. Hobart Bot. Gard. (T) 5-10 Inverted Prumnopitysferruginea Waitakere Hills (NZ) 1(-2) Inverted Prumnopitystaxifolia Manurewa Bot. Gard. (NZ) 5-10 Inverted a NZ = North Island, New Zealand, T = Tasmania, US = United States. b Morphological orientationwith respectto cone axis. consideredby Doyle to be highlyderived and Most observationswere made on populations analogousto Tsuga.Our observationsgreatly in theAuckland region of New Zealand (Table amplifythe picture for Podocarpaceae and show 1) sincethese provided convenient access to manyunique features of their pollination; per- photomicrographicfacilities. Pollination drops haps onlyin Picea and Pinus,which have an canbe observeddirectly in the field with a hand abilityto retainpollen prior to dropsecretion, lens. Detached shootswith ovulate cones at mayparallels be drawn(Owens, Simpson, and the stageof pollinationwere brought into a Molder,1981; Owens,Simpson, and Caron, shelteredroom, kept in water,and enclosedin 1987). a plasticbag. This usuallyresults in drop se- The Podocarpaceaeincludes 18 genera(17 cretionin a normalway. Pollination drop se- ifPhyllocladus is excludedas Phyllocladaceae; cretionis mostlyin thelate evening,and the Page, 1990) and about 200 speciesof shrubs dropsare normallyresorbed by late morning, and trees,with a widedistribution in theGon- but enclosedcut shootscan retaindrops for dwanaregion, i.e., presentday Africa,south- severaldays. In Microcachrystetragona, fresh eastAsia, Australasia, and SouthAmerica (De conesflown from Tasmania to Aucklandper- Laubenfels,1988). New Caledonia (18 spp.) mittedthe photographyof pollinationdrops andNew Zealand (17 spp).both represent local in thisway. Microphotography ofcones at the centersof diversity and endemism.Plants typ- stage of pollinationwas done with a Wild icallyoccupy montane habitats in thetropics, stereomicroscopeusing Ektachromecolor butin New Zealandare a majorcomponent of transparencyfilm (ASA 100);Figs. 1-9, 12-20 themixed podocarp forest at all altitudesand are black-and-whitereproductions from these are frequentlydominant. Recent generic re- transparencies. alignmenthas greatlyclarified the status of cer- Somemanipulation of ovules with and with- taingroups, chiefly by dismembermentof the outcaptured pollen grains was doneunder the largeand previously unnatural genera, Dacryd- microscopeto understandthe mechanicsof iumand Podocarpus (e.g., Tegner, 1967; Quinn, pollenmovement. Reorientation of ovules with 1970,1982, 1987; De Laubenfels,1972, 1988; enclosedpollen demonstrated the buoyancy of Page, 1989, 1990). pollenwithin natural drops, or withindrops simulatedby addingwater. MATERIALS AND METHODS RESULTS Observationswere made on field-growntrees fromwild and cultivatedpopulations in the Generalmechanisms- Cone orientation -An periodSeptember to December1989, in New intrinsicfeature of the pollination mechanism Zealand,Tasmania, and New Caledonia.Rec- of mostpodocarps is the erectorientation of ognitionof thepollination process was facili- thecone at thetime of pollination, commonly tatedby continualmonitoring of shootdevel- broughtabout by growth curvature
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