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Pacific Science (1979), vol. 33, no. 2 © 1980 by The University Press of . All rights reserved

Rhizophora in the 1

F. J. TAYLOR 2

ABSTRACT: stylosa Griff. is recorded from Moorea and Bora Bora in the Society Islands. Earlier records from the Society Islands of R. mangle L. by Forster (1786) and R. mucronata Lam. by Gray (1854) are probably the result of mislabeling, and there is no evidence that the present stands of Rhizophora are not recent introductions.

ON CAPTAIN COOK'S second voyage of dis­ repeated by Papy (1956: 189) in an account covery (1772-1774) the naturalists on board of the vegetation ofTahiti without, however, the Resolution were J. R. and G. Forster. any suggestion that he had seen the . The latter published a list of records In September 1975 I visited this locality and that included: "202 R. mangle, foliis acutis, confirmed the presence of fructibus sublatoclavatis M.S.V. p. 442 n. 5 Griff. Subsequently, I visited Bora Bora and Societatis, Amicorum, nouarum Hebridum recorded the same at Aheatauiti Bay, insulaeet nova Caledonia" (Forster 1786: 35). near Anau. While Rhizophora still occurs in the On Moorea, 63 trees were counted. Al­ Friendly Islands (Tonga), the New Hebrides, though most were about 2 m high, there were and , most botanists have well-grown trees up to 3 m high and numerous regarded the Society Island record as er­ seedlings about I year old. There was only roneous. As far as I can discover, Guillemin a narrow, discontinuous fringe of (1836-1837) was the last person to cite this along the shore, which can be correlated with record. In the first part of his paper he lists the low tidal range. (The mean range at R. mangle in his "Iiste des plantes qui existent spring tides is approximately 0.3 m.) The simultanement dans l'Archipel de la Societe Rhizophora occurred on both sides of the et dans les autres fIes de la Mer du Sud," bay, on coral sand to the west and on mud but Rhizophora is not mentioned in the sub­ and in depressions in grassy (Paspalum sequent enumeration of Tahitian plants. vaginatum Swartz.) areas to the east. As­ A second record of Rhizophora in the sociated with the trees was Acrostichum Society Islands, this time of R. mucronata aureum L. Fiddler crabs (Uca chlorophthala­ Lam., was made on the United States mus crassipes Adams & White) were present Exploring Expedition 1838-1842 (Gray in the substratum, with a mean of 35 crab 1854), but Gray considered the record to be holes/m 2 in the muddy areas and 5/m 2 in the result of mislabeling. the coral sand. Littorina scabra L., a common Indo-Pacific gastropod on , was found on the leaves. Behind the Rhizophora PRESENT DISTRIBUTION IN THE were scattered trees of Hibiscus tiliaceus L., SOCIETY ISLANDS which is the dominant shoreline species else­ where along the coast. In a guide to the Society Islands, T'Ser­ The Bora Bora locality was similar, but stevens (1950) noted the presence of man­ the trees were taller and all were growing groves at Vaianahe Bay, Moorea, a record on mud that supported a growth ofHalophila sp. and a population of fiddler crabs. There was no Acrostichum, but there is a small 1 Manuscript accepted 16 July 1978. 2 University of Auckland, Marine Research Labora­ fishing village to the rear of the site. There tory, R.D. I, Leigh, New Zealand. were no evident signs of clearing of the 173 174 PACIFIC SCIENCE, Volume 33, April 1979

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FIGURE l. The distribution of Rhizophora sty/osa (dotted line), based on Tomlinson (1978) and personal obser­ vations (1970). The broad arrows indicate the general direction of the south equatorial current. F, Fiji Islands; S, ; T, Society Islands.

Rhizophora by the villagers, though a number sonal observations 1970), which is 2400 km of the larger branches had obviously been to the west (Figure I). Although this distance cut with an axe. is not excessive for an ocean passage by a adapted to marine conditions, the ocean currents in this part of the Pacific Ocean travel westward, so that any natural DISCUSSION dispersal by floating would involve a con­ T'Serstevens (1950) assumed that the siderably longer voyage. Nor does R. stylosa Moorea Rhizophora population had been occur on the intermediate island group of deliberately planted. I have been unable to the , although there are ap­ trace any evidence for this. On Bora Bora parently suitable habitats. These are both the old local inhabitants declared that the arguments against a natural dispersal of the Rhizophora had been there "toujours." How­ species to the Society Islands. ever, there was a U.S. Naval Station at Anau If Forster's (1786) and Gray's (1854) re­ during World War II (from 1942 to 1946), cords of Rhizophora in the Society Islands and it is possible that it was then introduced were correct, it would raise the possibility accidentally from elsewhere in the Pacific. that it had been overlooked for 100 years. M. Erwin Christian examined the file on this It would also mean that the arrival of base for me, but found no references to Rhizophora there predated European dis­ mangroves. covery, though this would not necessarily The probability that Rhizophora stylosa imply natural dispersal as it could have been reached the Society Islands by natural dis­ introduced by early Polynesian voyagers as persal is low. The nearest locality is Samoa suggested by Chapman (1970) for R. mangle (where the plant is usually referred to as R. L. var. samoensis Hochr. in Fiji, Samoa, mucronata Lam., but this is the result of Tonga, and the New Hebrides. misidentification-Tomlinson 1978; per- Rhizophora is a Linnean genus, and Forster

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would interpret the species in terms ofSpecies in the Biological Faculty of the Lomonosar Plantarum (Linnaeus 1753). Here Linnaeus State University of Moscow. These presum­ recognized five species, of which four have ably come from Tonga, as the only record subsequently been transferred to other man­ for this species in Forster (1786) is "Namouka grove genera. The one remaining species is archipelagi, Amicorum." R. mangle. More recent taxonomists have Hou's statement is interesting in that the distinguished a number of other species, and specimen could not be located 100 years ago. in the latest revision, Hou (1960) recognized Seeman (1865-1873: 92) comments: "Forster seven species, five of which occur in the mentions a mangrove as occurring in the Pacific. Though there is no general agreement Society Islands, but there is no specimen from on the status of several of the taxa, this is there." It may be that Seeman did not consult irrelevant to the present argument. any plants at Kew, though this seems un­ Forster's record of likely, for although he worked and consulted could therefore have been of another taxon, Forster's herbarium in the British Museum, possibly R. stylosa, and this is borne out by he deposited his specimens at Kew. his mention of "foliis acutis," which is There is another possible line of attack characteristic of the Pacific taxa other than on the identification. Hou mentions that R. mangle [and the Fijian/Samoan/Tongan/ Forster's specimen bears the annotation New Caledonian taxon variously included "common name: wabitatin malabar." The in R. mangle L. or R. samoensis (Hochr.) linguistic origin of this name appears to be Salv., but which Tomlinson and Womersley Melanesian rather than Polynesian. The (1976) recognize as R. mangle var. samoensis general Polynesian term for mangroves is Hochr.]. Rhizophora mangle and R. samoensis tonga (R. M. Clark in litt. 1977) [cf. togolei have obtuse leaves. That Forster included given by Yuncker (1959) as the local Tongan these other species in his concept of R. mangle name for R. samoensis]. This suggests that is made clear by his record of it from the the specimen was not Polynesian, and thus New Hebrides (Forster 1786). In his account not Tahitian. Unfortunately, the limited ofthe voyage (Forster 1777) the only mention amount ofmaterial available on the languages of mangroves in the New Hebrides comes of New Caledonia and the New Hebrides from Port Resolution on the southeastern [the only Melanesian localities given by corner of Malekula Island. The only Rhi­ Forster (1786) for R. mangle] does not permit zophora species present at Port Resolution a more definite localization at present. are R. stylosa and R. apiculata Blume (per­ In his account ofthe voyage Forster (1777) sonalobservations, 1974). mentioned mangroves in Tonga, New Cale­ These problems could be cleared up by donia, and the New Hebrides, corresponding examination of Forster's specimen. Hou with his published records (Forster 1786), (1960) states that it is at Kew, and he ex­ but he does not mention them while in the amined it there (c. Kalkman in litt. 1977), Society Islands. M. E. Hoare (in litt. 1977) but searches of the Pacific Rhizophora there examined Forster's unpublished journal and have failed to reveal it (P. S. Green in litt. has confirmed that there is no mention there 1976; M. Bywater in litt. 1977). Correspon­ of mangroves in the Society Islands. On dence with the herbaria indicated as having , Forster botanized mainly around Forster material in Stafleu and Cowan (1976) Point Venus. Mangroves do not grow there has failed to bring any of his Rhizophora today and the shore there is not suitable for material to light. The only Forster mangrove them (personal observations 1975). He did material it revealed was a sheet of leaves not visit either Moorea or Bora Bora, but labeled "Rhizophora gymnorhiza" [Brugiera did visit , which I was unable to visit. gymnorhiza (L.) Lamk.] without locality or and R. stylosa have date, in the Systematisch-Geobotanisches been confused for many years, and Gray's Institut, G6ttingen, and a single leaf with (1854) record of the former could well be the same annotation and no date or locality of the latter. Unfortunately, the specimen

, . 176 PACIFIC SCIENCE, Volume 33, April 1979

was described as "a fragment," and again Cook, during the years 1772, 3, 4 and 5. it has not been located in the U.S. National London. 2 vols. Herbarium at the Smithsonian Institution ---. 1786. Florulae insularum australium (J. J. White in litt. 1977), so that it is not podromus. Gotting. 103 pp. possible to check the identification. Gray also GRAY, A. 1854. United States Exploring notes that in his narrative of the voyage, Expedition during the years 1838, 1839, Pickering does not mention mangroves in 1840, 1841, 1842 under the command of the Society Islands. Charles Wilkes, U. S. N., Botany. Phanero­ The two early records lead one to think gamia. Vol. 1. that Rhizophora slylosa possibly did occur GUILLEMIN, J. B. A. 1836-1837. Zephyritis in the Society Islands 150-200 years ago and Taitensis. Enumeration des plantes decou­ has been overlooked since. The very low vertes par les voyageurs dans les lIes de (0.3 m) tidal range results in discontinuous la Societe, principalement dans celie de stands of Rhizophora that could easily be Taiti. Ann. Sci. Nat. Ser. 2, 6:297-320; missed by casual observers making short 7:177-192,241-255,349-370. visits to the more remote islands. However, Hou, D. 1960. A review of the genus Rhizo­ the absence of a mention of mangroves in phora with special reference to the Pacific the expedition journals suggests that the species. Blumea 10: 625-634. evidence is in favor of the mislabeling of the LINNAEUS, C. 1753. Species plantarum. 2 vols. earlier specimens, though if there were only Stockholm. a few trees present, they might not merit a PAPY, H. R. 1956. Tahiti et les iles voisines. mention. There is thus no reason to suggest 2e partie. La vegetation des iles de la that the present stands are not recent intro­ Societe et de Makatea. Trav. Lab. Forest. ductions, though they already appear to have Toulouse. V. Sect. 2. I(III): 163-386. developed part of the characteristic man­ SEEMAN, B. 1865-1873. Flora Vitiensis. A grove fauna. description of the plants of the Viti or Fiji Islands. London. 453 pp. and atlas. STAFLEU, F. A., and R. S. COWAN. 1976. Taxonomic literature. Vol. 1. 2d ed. Bohn, ACKNOWLEDGMENTS Scheltema, & Holkema, Utrecht. I am grateful to all the correspondents who TOMLINSON, P. B. 1978. Rhizophora in Aus­ have replied to my queries, to the directors tralasia-some clarification of and staff of the various herbaria, and to P. B. and distribution. J. Arnold Arbor. 59: Tomlinson for a draft of his paper on 156-169. Rhizophora. TOMLINSON, P. B., and J. S. WOMERSLEY. 1976. A species of Rhizophora new to New Guinea and , with notes relevant to the genus. Contrib. Herb. Aust. LITERATURE CITED 19: 1-10. CHAPMAN, V. J. 1970. Mangrove phytosoci­ T'SERSTEVENS, A. 1950. Tahiti et sa Cou­ ology. Trop. Ecol. 2: 1-19. ronne. I. Tahiti, Moorea; les Polynesiens. FORSTER, G. 1777. A voyage round the Michel, Paris. world in His Britannic Majesty's Sloop, YUNCKER, T. G. 1959. Plants of Tonga. Resolution, commanded by Capt. James Bernice P. Bishop Mus. Bull. 220: 1-283.

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