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Seed Morphology and Anatomy of Rhizophoraceae, Inter-and Infrafamilial Relationships

Hiroshi Tobe and Peter H. Raven
Annals of the Missouri Botanical Garden
Vol. 75, No. 4 (1988), pp. 1319-1342
DOI: 10.2307/2399288
Stable URL: http://www.jstor.org/stable/2399288
Page Count: 24
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Seed Morphology and Anatomy of Rhizophoraceae, Inter-and Infrafamilial Relationships
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Abstract

We present an overall study of the seed morphology and anatomy of all Rhizophoraceae (10 inland and 4 mangrove genera). Morphologically seeds are arillate, winged or nonappendaged; both arillate and winged seeds are borne in capsular fruits, and nonappendaged seeds in baccate or indehiscent hard-walled capsules. Seed coat anatomy is diversified in correlation with the seed and fruit morphology, but a well-developed exotesta and a fibrous exotegmen are common to all inland genera. Despite certain minor divergences, the seed coat of different genera of Rhizophoraceae is defined as exotestal, exotestal-exotegmic, or undifferentiated. Different combinations of seed morphological and anatomical features characterize different genera or groups of genera. An overall comparison of seeds and other reproductive characters confirms that in Rhizophoraceae arillate seeds and the presence of a fibrous exotegmen are plesiomorphic features that can be used in searching for related families. Seed morphology and anatomy also support grouping Rhizophoraceae with Elaeocarpaceae and Celastraceae, and the exclusion of Elaeocarpaceae from Malvales. Our comparison further suggests that the seed morphology and anatomy of Rhizophoraceae have evolved as the result of adaptation to different methods of seed dispersal from ant dispersal to dispersal by wind, bird, mammal, or water. We carried out a cladistic analysis of the genera of Rhizophoraceae on the basis of 16 reproductive characters (including those of seed morphology and anatomy). Blepharistemma, Cassipourea, Comiphyton, Anopyxis, Macarisia, and Sterigmapetalum, all inland genera that have been assigned to Macarisieae, are characterized by having many plesiomorphies (particularly a superior ovary). The last three genera have winged seeds and a thinner seed coat (apomorphies), whereas the first three have arillate seeds. Among the four remaining inland genera, which have been assigned to Gynotrocheae, Crossostylis (with arillate seeds) differs greatly from the others in having only one apomorphy (i.e., an inferior ovary); it retains many plesiomorphies. In contrast, Carallia, Gynotroches, and Pellacalyx share a nonappendaged seed and a persistent meso- and endotegmen, both clearly synapomorphies, and Gynotroches and Pellacalyx further share some distinct synapomorphies, suggesting their close affinities. The four mangrove genera-Bruguiera, Ceriops, Kandelia, and Rhizophora-which have been segregated as Rhizophoreae, share many synapomorphies (e.g., an undifferentiated seed coat and the complete lack of the tegmen), and therefore the coherence of Rhizophoreae is not in doubt. Cladistically the mangrove genera are more closely related to Carallia, Gynotroches, and Pellacalyx than to Macarisieae. Based on our cladistic analysis, the traditional infrafamilial classification is revised, and a new tribe Crossostylideae, which consists of Crossostylis only, is proposed.

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