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Preliminary Observations on Floral Biology in Mangrove Rhizophoraceae Author(S): P

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Preliminary Observations on Floral Biology in Mangrove Rhizophoraceae Author(S): P Preliminary Observations on Floral Biology in Mangrove Rhizophoraceae Author(s): P. B. Tomlinson, R. B. Primack, J. S. Bunt Source: Biotropica, Vol. 11, No. 4 (Dec., 1979), pp. 256-277 Published by: The Association for Tropical Biology and Conservation Stable URL: http://www.jstor.org/stable/2387918 . Accessed: 30/08/2011 15:41 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 Association for Tropical Biology and Conservation is collaborating with JSTOR to digitize, preserve and extend access to Biotropica. http://www.jstor.org PreliminaryObservations on Floral Biology in Mangrove Rhizophoraceae P. B. Tomlinson, Harvard University,Harvard Forest, Petersham, Massachusetts 01366, U.S.A. R. B. Primackt Department of Botany, Universityof Canterbury, Christchurch, New Zealand and J. S. Bunt Australian Instituteof Marine Science, Townsville, Queensland, Australia ABSTRACT The tribe Rhizophoreae (Rhizophoraceae) includes the largesttaxonomic assemblage exclusivelyof mangroves,with 4 genera and about 18 species. Rhizophora is pan-tropical;Bruguiera, Ceriops, and Kandelia have an Indo-Malayan dis- tribution.All species have the same basic floral structure,but field observationsdemonstrate a wide varietyof pollina- tion mechanisms.Variation in such featuresas size and orientationof flowers,number of flowersper inflorescence,num- ber of stamens,time of stamendehiscence, and method of pollen dischargecan be shown to have direct relevance to pol- lination biology. Pollen vectorsmay be predominantlywind (Rhizophora, as deduced from circumstantialevidence) or animals (remaining genera, as observed directly). An explosive method of pollen release occurs in Bruguiera and Ceri- ops tagal, but is modifiedto suit a range of flowervisitors such as birds in the large-floweredBruguiera species (e.g., B. gymnorrhiza)or butterfliesand other insectsin the small-floweredBruguiera species (e.g., B. parviflora) and moths in Ceriops tagal. Ceriops decandra lacks the explosive mechanismof pollen release,as does Kandelia. Since these taxa often formmixed communitiesa high degree of partitioningof pollinator resource is achieved. INTEREST IN T HE REPRODUCTIVE BIOLOGY of man- mentalstudies of fluid-preservedmaterial, which will groves has largelycentered on seedling development form the basis for a separate comparative account. and dispersal since embryodevelopment on the par- All taxa in the section Rhizophoreae of Rhizophor- ent plant to a seedling stage (vivipary) is more fre- aceae have been studied, and the comparative data quently shown in this group of plants than in any accumulated permit some speculation about floral other. Studies on floral ecology are very few, as van evolution,although this must remain tentativeuntil Steenis (1958) points out, but knowledge of repro- comparable informationfor non-mangrove Rhizo- ductive capacity and interspecificsterility barriers phoraceae has been assembled. must begin with a considerationof flower function. That the Rhizophoreae include distinctivefloral The most extensive studies are those of Guppy mechanismsis evident from the early summaryand (1906) on Rhizophora. The present account is a reportsof Gehrmann (1911) and Porsch (1924, p. summaryof informationobtained during field work 611) who recorded the explosive method of pollen in mangrove communities,mainly in Papua New release in Bruguiera sexanzgula,associated with bird Guinea and Queensland, and surveys floral biology visitors,an observationmore recentlyconfirmed by of Rhizophoraceae in a provisional way. Field work Davey (1975) in southeastAfrica. We can now re- provided the opportunityto observe flower visitors port that all Bruguiera species have explosive pollen and floral mechanisms in Bruguiera, Ceriops, and release,but that two main agents triggerthe process, Rhizophora. Additional informationhas come from insects in the small-floweredspecies, birds in the other parts of the South Pacific (mainly Fiji and large-floweredspecies. Ceriops tagal is similarlyex- New Caledonia) and in South Florida. No field ob- plosive, but triggeredmainly by moths. Ceriops de- servationson Kandelia have been made. Although candra lacks explosive dehiscence,and pollen vectors informationis incomplete it is sufficientto dem- are as yet unknown. onstrate a wide range of pollination mechanisms Despite the abundance and wide distributionof which occur in plants with a fundamentallysimilar Rhizophora there is no extended treatmentof its floral construction,suggesting a fairlycomplete di- floral biology. Guppy (1906) provided information vision of pollinator resources. about floral mechanism in Fijian Rhizophora spe- The field observationshave been supplemented cies, but did not commenton vectors.Gill and Tom- by detailed morphological,anatomical, and develop- linson (1969) have suggested, but without much supportingevidence, that R. mangle is wind-pollin- lPresent address: Department of Biology, Boston Univer- sity,Boston, Massachusetts02215, U.S.A. ated; Kress (1975) observed insect visitors to Rhi- 256 BIOTROPICA 11(4): 256-277 1979 zophora in South Florida; and there is one recordof unopened flowers.The antherswere maceratedcom- bees visiting the flowersof R. stylosain Java (Doc- pletely in several drops of filteredsea water. Addi- ters van Leeuwen, 1927). Our comparativeobserva- tional filtered sea water was added to bring the tions lead us to the conclusion that the flowersof suspensionof particulatesup to a volume of approxi- most Rhizophora species are basically wind-pollin- mately 10 ml. The suspensionwas violentlyagitated ated, although they may be a pollen source for in- for 30 seconds to freeany pollen grains still enclosed sects and may even retain vestiges of insect attrac- by the anther tissue, and then filteredthrough a 50 tion in odor and nectar. Rhizophora stylosa appears micronscreen mesh to remove tissue debris.Virtually to be pollinated by small bees and wind. no pollen grains were trapped in this screen,nor did Since the taxa involved are wide-ranging (pan- theyclump or settle,at least within the few minutes tropical in Rhizophora) and our field work is ne- that the suspension was utilized. A Coulter Counter cessarilygeographically restricted, it is clear that our (model TAII, Coulter Electronics,Hialeah, Florida), descriptionsmust serve only as an introductionand normallyused for countingcell numbersas in blood our conclusions may not be universallyapplicable. samples, was then used to count the numberof pol- Further field studies in differentgeographic areas len grains in a subsample of this suspension,or if are needed. At the same time our observationsare the pollen densitywas still too high to count, in a sufficientlycomplete to indicate an example of adap- more dilute suspension. Successive subsamples from tive biological radiation in a widespread and eco- the same pollen suspension gave virtuallyidentical nomically importanttropical group which is prob- counts.The numberof pollen grains in each of two ably monophyletic. anthers of Bruguiera parviflora,Ceriops tagal, and Kandelia candel were counted completelyafter the antherswere maceratedin a drop of iodine solution. MATERIALS AND METHODS In order to confirm the results of the Coulter Most detailed observationswere made at field sites Counter,two antherseach of Bruguieragymnorrhiza, in Queensland (3 Mile Creek and Cape Ferguson, Rhizophorazmucronata, R. X lamarckii,and R. apicu- Townsville and Missionary Bay, Hinchinbrook Is- lata were macerated,agitated and suspended in 4 ml land), but these have been supplemented by more of a solution of 1 part aqueous iodine to 2 parts gly- general observationsin Australasia.Unless otherwise cerine. The pollen grains suspended in one drop stated,observations refer to the Queensland localities. (0.025 ml) of this solutionwere counted.This man- ual countingmethod gave similarresults to the Coul- ter Counter figures,supporting the validity of both TABLE 1. The numberof pollen grainsper flower(P), approaches.The resultsare presentedin table 1. ovulesper flower(0), and pollengrains pro- ducedper ovule (P/O) in speciesof the Rhi- zophoraceae.The numberof pollengrains per STIGMA RECEPTIVITY.-The onset of stigma recep- flowerand per ovule are given in thousands; ovule numbersare the most commonlyob- tivityin Bruguiera exaristataand Rhizophora stylosa servedvalues. was determinedby floating stigmas in a naphthyl- acetate. This produced a color reaction, i.e., meta- Pollen Genus and species P x 103 0 P/O x 103 vector bolically active surface cells indicative of esterase activity,by coupling with hexazotised pararosanilin, Rhizophoraapiculata 5256 4 1314 R. mucronata 2096 4 524 Wind which in turn is indicative of stigma receptivity R. X lamarckiia 4000 4 1000 (Heslop-Harrison et al. 1975). Bruguiera (Type 1) B. gymnorrhiza 1800 6 300 B. exaristata 1080 6 180 Birds TAXONOMY Bruguiera (Type 2) B. cylindrica 176 6 29 The section Rhizophoreae of Rhizophoraceae in- B. parviflora 4.8 6 0.8 Insects Ceriops decandra 84 4 21 (Insects?) cludes four genera distinguishedby their exclusively Ceriops tagal 9.2 6 1.5 Moths mangrovehabitat
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