mangrove-associated algae in full
TRANSCRIPT
MANGROVE-ASSOCIATED ALGAE IN FUll
ABSTRACT
R. Ashni Raj Prasad and G. Robin South
Marine Studies Programme The University of the South Pacific
Suva Republic of Fiji
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Macroscopic algae epiphytic on mangrove pneumatophores were surveyed from two climatically contrasting mangrove communities in the Ba and Rewa River mangrove swamps on Viti Levu, Fiji. Seventeen taxa, including 13 Rhodophyta, 2 Phaeophyta and 2 Chlorophyta were recorded. The most frequently occurring species is Dictyota bartayresiana, followed by Chlorodesmis hildenbrandtii and Caloglossa leprieurii. Two genera normally associated with mangroves in other regions were absent in Fiji: Catenella and Murrayella. Compared with other studies in the Indo-West Pacific region, Fiji has the lowest number of mangrove-associated algae, and a greatly reduced number of mangrove species. On a localized scale, however, variations in climatic conditions (between Ba and Rewa) do not seem to affect the associated algal epifiora, and no mangrove host-specificity is apparent.
INTRODUCTION
Mangroves reach their maximum development and greatest luxuriance in Southeast Asia (Clayton and King, 1991), and the best developed mangal is found near the equator in Indonesia, Papua New Guinea and the Philippines. Species numbers decline with increasing latitude, and with distance from the Indo-Malaysian region (Chapman, 1984). Mangroves do not occupy large areas in the Pacific Islands. Of the approximately thirty species in Papua New
'Dedicated to the Memory of William R. Kenchington and in recognition of his lifetime contributions to The University of the South Pacific and to the South Pacific Journal of Natural Science.
S.PacJ.Nat.Sci., 1995,14:115-131
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Guinea and Australia, nineteen extend to the Solomon Islands, eleven to Vanuatu and New Caledonia and 8 reach Fiji and Tonga (Wood roffe, 1987). The eastern limit of mangrove forests in the Central Pacific is in Annu 'u in Samoa, with 4 species (Wood roffe, 1987). Woodroffe (1987) also reports on species numbers in other Pacific Islands as: Micronesia (1 0), Marshall Islands and Kiribati (4) and Tuvalu (2). Mangroves are absent from the Kermadecs, Niue, Cook Islands, Tokelau and the Phoenix Islands (Kiribati), and they are considered to have been introduced to Tahiti, the Society Islands, Eniwetok and Biki ni (Woodroffe, 1987).
The Fijian mangrove flora includes the following species: Bruguiera gymnorhiza (L.) Lam.; Rhizophora samoellSis (Hochr.) Salvoza; Rhizophora X selala (Salvoza) Tomlinson (a putative hybrid of R. samoellSis and R. stylosa); Rhizophora stylosa Griff.; Xylocarpus granatum Koenig; Lumnitzera litlorea (Jack.) Voigt; Excoecaria agallocha L; and Heritiera littoralis Dryand (pillai, 1988). Four taxa dominate the flora:B. gymnorhiza, R. stylosa, R. samoellSis and the hybrid R. x selala. The flora has been reasonably well studied (parham, 1972; Tomlinson, 1978; Raj and Seeto, 1982; Raj. et aI., 1984; Watling 1985,1986; Pill ai, 1984, 1988).
Mangroves play an important role in coastal ecology, and have an accompanying specialized epiflora, the most successful of which clothe the branches of the trees up to their extreme tips (Saraya, 1985). The epiphytic community consists of macro- and microalgae, mosses, ferns, lichens and flowering plants of tropical and subtropical regions. They are only attached to, and are not parasites on mangroves (Saraya, 1985). Oliviera (1984) divides algae found in the mangal into two main groups: the planktonic algae (especially diatoms) and the benthic algae, which he further divides into those occurring on trunks, prop roots and pneumatophores of mangroves, and those that occur on soil and rock.
The most detailed earlier accounts of mangrove-associated algae are those of Post (1936,1963,1967, 1968). A numberof recent studies have applied modem taxonomic treatments to mangrove-associated algae from various parts of the world, including South Australia (Beanland & Woelkering, 1982, 1983); Victoria (Davey & Woelkerling, 1980); New South Wales (King & Wheeler, 1985); Indonesia (Tanaka & Chihara, 1988a, b); Japan (Tanaka & Chihara, 1984a, b); South Africa (Lambertet al., 1987); Papua New Guinea (King, 1990); Kenya (Coppejans & Gallin, 1989); Brunei, Darussalem (King et al., 1991b) and North America (Sheath et al., 1993). In addition, detailed studies of the
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taxonomy of mangrove algae have greatly improved our knowledge of the component species (King & Puttock, 1989, 1991; Kinget al., 1988; Sheath et
ai.,1993).
Very little has been published on mangrove algae in Fiji: reference is restricted to some brief comments in Troll (1931), Post (1936,1967), Chapman (1976) and Hono (1985). South & Kasahara (1992) include previously reported mangroveassociated algae in their preliminary checklist..
In this account we describe the results of a floristic survey of mangrove associated macroscopic algae made in March - September 1992 at two climatically contrasting sites, the Ba and Rewa Rivers on the island of Viti Levu (Fig. 1). Ba lies in the rain shadow part of the island, while the Rewa delta lies on the wet, windward side. The study attempts a first review of the Fijian mangrove-associated algal flora, while at the same time aims to compare the epiflora between the two contrasting sites, and the Fijian flora with that of other regions in the Indo-West Pacific. In addition, an analysis of the flora in relation to mangrove host species is provided.
MATERIAIB AND MEfHODS
The Ba and Rewa sites (Fig. lA, B) were selccted from aerial photographs and topographic maps, and a thorough study was made of previous surveys carried out by Watling (1985). A general comparison of the climate at the two sites is presented in Table 1.
The Study Sites - The Ba Delta (Fig. lA)
The Ba Delta includes an area of c. 3,995 ha of mangroves (Watling, 1985). The relatively dry, seasonal climate (Table 1) has a profound effect on the mangrove vegetation, especially in the more elevated and less well-drained areas that are poorly flushed by high tides and so become hypersaline.
Floristically, the Ba mangrove is the least diverse, with two major mangrove alliances distinguished: the Tiri and Selala Alliances (Watling, 1985). There is a complete absence of a Bruguiera gymnorhiza dominated forest.
In the Tiri Alliance (transects 1-5 and 10-16, Fig. 1A), Rhizophora stylosa forms an almosfpure closed shrub forest on the extensive poorly drained flats behind river and creek beds, and the hypersaline flats are surrounded by stunted forms (Watling, 1985).
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Figure 1 • A, B Map of Viti Levu, Fiji, showing the location of the Ba (A) and Rewa (B) deltas. Locations of the 16 transects at each site are shown in black: for details seetext.
50km
The Selala Alliance (transects 6 - 9, Fig. lA) is an open forest alliancedominated by Rhizophora xseLala, found to a varying extent on almost all theriver and creek banks and associated water course levees (Watling, 1985).Bruguiera gymnorhiza and Xylocarpus granatum occur infrequently witheither R. styLosa or, more commonly, withR. samoensis.
The Rewa Delta (Fig. lB)
The Rewa Delta is on the windward side of Viti Levu, experiencing high andmore frequent rainfall , higher humidity and more even temperatures (Table I). The extent of disturbance caused by cyclones is less than on the rain shadowside. The Rewa Delta is the most fertile and productive in Fiji, supportingabout 5,130 ha of mangro~es . The mangrove stands are found in the backwaters, protected from strong wave action by the Suva Barrier Reef and other
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associated reef systems. The mangrove associations at Rewa are the most diverse in Fiji, and six alliances have been distinguished (Watling, 1985): the Seaward Alliance; Tiri Alliance, Dogo Forest Alliance, Mixed Alliance, Landward Alliance and the Boreti Alliance. Rewa transects were made in the Seaward, Tiri (transect 11 only), Dogo Forest and Landward Alliances (Fig.IB).
The Landward Alliance (transects 1-8, Fig. IB) is an heterogenous closed forest which forms the floristically most diverse of Fiji 's mangrove alliances. It is found only in the least exposed situations, with extensive epiphytic and climbing flora. The most common mangrove species are Bruguiera gymnorhiza, Xylocarpus granatum and Excoecaria agallocha. Rhizophora samoellSis and R. x selala occur more rarely and on creek banks. Other common trees of this alliance are given in Watling (1985).
TheDogo Forest Alliance (transects 9-10; 12-14, Fig.lB) isa uniform closed forest of almost pure G. gymnorhiza, with occasional Rhizophora x selala, XyLocarpus granatum and Lumnitzera littorea. In many instances, the landward Alliance and Mixed Alliance merge into this Alliance (Watling, 1985).
The Seaward Alliance (transects 15, 16) is a mixed fringing forest, found usually on the more exposed seaward banks. Rhizophora stylosa often forms a single belt on the seaward edge with taller trees abruptly behind, mixed with Rhizophora x selala, Bruguiera gymnorhiza and R. samoellSis. Xylocarpus granatum and Excoecaria agallocha are found in the landward region (Watling, ]985).
Sampling methods:
A summary of the 16 randomly selected transects made at each site is presented in Table 2. Along each transect, five pneumatophorcs, prop roots or knee roots (depending on the host species; they are collectively referred to here as pneumatophores) closest to the midpoint of each quarter of the transect length were collected and placed in labelled plastic bags. All macroscopic algae epiphytic on the pneumatophores were included in the analysis. Mangrove species were identified and a listing of all algae occurring on the 310 pneumatophores for 16 transects at each of the two localities formed the basis of the analysis.
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Analysis of samples:
On return to the laboratory, all algae were carefully scraped from whole pneumatophores and preserved in 5% formaldehyde in sea water for later examination and identification. Representative specimens are preserved in the Phycological Herbarium, South Pacific Regional Herbarium (USP) as slides or herbarium specimens.
The total number of species was recorded, according to site (Ba and Rewa) and host mangrove. Frequency of each species at each locality was calculated using the formula in Beanland & Woelkering (1982), and as used in King & Wheeler (1985) and Davey & Woelkerling (1985):
F = S N/N X 100, where
F = percentage frequency; SN = the number of pneumatophores on which a particular species occurred, and N = the number of pneumatophores surveyed.
For ease of comparison with previously published data from similar studies, the "relative profusion of taxa" was determined (Bean land & Woelkering, 1982; Davey & Woelkerling, 1985). Species have been assigned to one of the five categories: rare (F <.5%), sporadic (F ;;:;: 5-24 %); occasional (F = 24-49%); common (F = 50-75%); and abundant (F >75%).
Mean percentage frequency (F) data for each species occurring at the two sites were calculated using the formula in King & Wheeler (1985):
F = Ft/N X N, where
F;;:;: mean percentage frequency; Ft = sum of all recorded frequencies >0; N = the total number of localities at which the alga occurred. The relationship between the number of common species (F >50%) and the total ~umber of species found at the two sites was calculated using the method of Davey &
. Woelkerling(1980):
Ps/Ts, where
Ps = the number of common species (F > 50%); Ts = the total number of species.
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Observations of possible host specificity of algal species were made for both localities. In addition, a comparison was made between Ba and Rewa for the total number of algal species found per host mangrove species.
RESULTS
A total of 17 taxa of mangrove-associated macroalgae was recorded (Table 3). Descriptions, taxonomic and nomenclatural notes on the component species are given in Raj (1993) and South et al. (1993). At both sites, 16 ofthe 17 taxa were found.
Frequency data are shown in Table 3. Four taxa occurred commonly [three in Ba and one in Rewa: CalogLossa leprieurii (Ba),Rhodolachne radicosa (Ba), Dictyota bartayresiana (Ba) and ChLorodesmis hildebrandtii (Rewa)]. Based on mean frequency values, Dictyota bartayresiana is the most frequently occurring species (F=40), followed by Chlorodesmis hiLdebrandtii (F=38), Caloglossa leprieurii (F=36),Bostrychia radicans and RhodoLachne radicosa sp. inedit (F=35). The least frequently occurring was Polysiphonia sp. (F=3). Except for Polysiphonia sp. and Bostrychia tenella, all other taxa occurred at both sites. None of the taxa was abundant (F> 75).
Table 4 provides a summary of the algal species found for the different host mangrove species sampled at Ba and Rewa. No host specificity is shown by any of the species, since they occurred on all hosts at either of the two sites, if not at both sites. There was a difference in the mean number of algal taxa occurring per host tree between the two sites, with 13.6 at Ba, and 15.3 at Rewa.
DISCUSSION
While our study was restricted to two contrasting sites, and may not represent an exhaustive list of species, there is strong evidence to suggest that the decline in host species diversity in the Indo-Pacific region is paralleled by a decline in the diversity of the associated macroalgal flora. As for other mangrove algal floras, the Rhodophyta were dominant, but in Fiji two genera commonly found in the mangrove flora elsewhere, Catenella and MurrayeLLa, were absent, while the number of species recorded was comparatively low. The different host species diversity at the two climatically contrasting sites did not appear to influence the numbers of algal species present, and there was no evident host specificity.
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For all studies the Rhodophyta are dominant. King & Wheeler (1985) showed in their study of mangrove macroalgal communities in New South Wales, Australia that the most frequently occurring algae were those of the Bostryehia - Caloglossa association. Davey & Woelkerling (1980) in their studies of mangrove algae in Victoria Austral ia showed that the Victoria flora is comparable in species numbers with that of Queensland, but that the species composition of the two areas differs markedly, with none of the Chrysophyta or Phaeophyta common to both regions. Comparison of the New South Wales macroalgal mangrove flora with that of eastern and southern Australia shows greatest similarity in both number of species and shared species, with that of southern Queensland (King & Wheeler, 1985).
Mosisch (1993) has studied the effects of salinity on the distribution of two populations of CaLogLossa Leprieurii in the Brisbane River, using transplant experiments and laboratory culture. Each showed different salinity tolerance ranges and corresponding differences in distribution, indicating that there are distinct salinity ecotypes.
Lambert et al. (1987) investigated macroalgae associated with 19 mangrove estuaries in South Africa and observed that persistent components of the flora are Bostryehia tenella, B. radieans and CalogLossa leprieurii. In the eastern Indonesian mangrove forests, species of Bostryehia, CaLogLossa, and CaleneLLa are dominant (Tanaka & Chihara, 1988a, b). A similar situation is reported from Brunei Darussalam (King et at., 1991) and for Papua New Guinea (King, 1990).
CateneLla and MurrayeLLa are apparently absent from Fiji, although both occur in Indonesia (Tanaka & Chihara 1988a, b) and Papua New Guinea (King, 1990). In Fiji, the most commonly occurring species were CalogLossa leprieuri4 RhodoLaehne radieosa (= Bos/ryehia sp.; see Table 2), Dietyota bartayresiana, and ChLorodesmis hildebrandtii (Rewa). In New South Wales King & Wheeler (1985) recorded Bostryehia moritzialla, B. simpliciuseula and CalogLossa Leprieurii as the most common or abundant (F=50-75%, or F> 75%). For the Victorian flora, Davey & Woelkerling (1985) indicated that CalogLossa lepieurii occurred most commonly (F=50-75%).
Bostryehia simplieiuscula has not been previously recorded from Fiji (South et al., 1993); King & Puttock (1989, 1991 a) have provided detailed taxonomic studies of this species. Two of the Caloglossa species are to be re-named (King'& Puttock, pers. comm.), while Gelidium erinale var. perpusillum is
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also new for Fiji (South et al., 1993); it was reported from Indonesian mangroves by Tanaka & Chihara (1988a, b). Rhodolachne radicosa, described by ltono (1985) appears to be a species of Bostrychia (Puttock, pers. comm.).
The more diverse mangrove-associated algal flora closer to the centre of distribution of mangroves in the Indo-West Pacific region may be due to a wider range of available habitat. Past information has suggested, however, that like the mangrove species, there is in the Pacific Islands a general reduction in the numbers of macroalgae from west to east (Payri, 1991). This is, however, partly a reflection of the lack of detailed floristic studies in isolated Pacific Islands; the Fijian algal flora, for example, is now known to be much more diverse than previously recognized, and it is estimated that only 60 - 70% of the flora is currently reported (South et aI., 1993).
Unlike other elements of the flora, the mangrove-associated algae are a special case, largely depending on the microhabitats made available by their hosts. While it is possible that more extensive sampling might reveal more mangrove-associated species in Fiji, the relative paucity of species is likely to remain, a reflection of the comparatively small numbers of host species.
This investigation generally underscores the relative uniformity of the mangrove-associated algal flora world-wide. There is a parallel trend in the host and associated species impoverishment from west to east in the IndoPacific, and the disjunct distribution of mangroves on increasingly isolated islands and island groups is paralleled by the composition of the associated algal flora. Studies on other island groups are likely to confirm this; for example, mangroves found on the Tarawa Atoll, Kiribati, are largely devoid of algal epiphytes (South, unpubl.). The fact that distinct populations of mangrove-associated algae appear not to have evolved on isolated islands suggests that their arrival may be relatively recent, and that they have travelled together with their hosts.
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ACKNOWLEDGEMENTS
This work is part of the thesis submitted by RARP for the M.Sc. degree at the University of the South Pacific. We are grateful for financial support from the University of the South Pacific (RARP) and from the International Centre for Ocean Development, the Canadian International Development Agency, and the Government of France (GRS). We thank Assoc. Prof. Robert J. King and Dr Christopher F. Puttock for valuable assistance with identification of species. The study would not have been possible without the field assistance of Dharamraj and Mukesh Raj, and friends Praveen and Pradecp.
TABLE 1
Summary of climate in Ba (upper line) and Rewa (Iowerline). Ba information based on data for Nadi (Sharma, 1982); Suva information based on © PC Globe, Inc. (1992) and Morrison & Naqasima (1992).
Parameter F M A M A S 0 N
Mean 298 286 353 180 84 70 49 59 87 98 141
rainfall 290 272 368 310 257 170 124 21 196 21 1 249
(mm)
Mean Temp. 27 26 26 26 25 24 23 24 25 25 26
(OC) 27 27 27 26 25 24 23 23 24 24 25
ReI. Humid. 79 80 82 81 79 78 72 70 72 72 73
(%) 85 85 85 85 83 83 81 81 82 82 82
0
17 1 31 -
27
261
75 ' 83
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TABLE 2 Details of transects, Ba and Rewa
Transect Date No. pneum. Transect Mangroves (,92) length(m)
BA 1 2MB :!) 20 1,3,4,5,6
2 22;U3 :!) 25 3,4
3 22;U3 :!) 25 3
4 03~ :!) 25 4,6
5 03~ :!) 100 3,4,6
6 ~ :!) 25 4,5,6
7 OOft)S :!) 35 3,6
8 1QUS :!) 100 4,5
9 1QUS :!) 16 4
10 23ft)S :!) 10 4
11 23ft)S :!) 10 1,3,4
12 24ft)S :!) 100 4
13 24ft)S :!) ]00 4
14 25m :!) 100 3,4
]5 2fJI(J7 :!) ]00 3,4
]6 2fJI(J7 :!) ]00 4,5
REWA 1 <X>tV4 :!) 28 1,4
2 <X>tV4 :!) 44 1
3 <X>tV4 :!) 60 1,3
4 <X>tV4 :!) 60 6
5 17~ :!) 60 1,3
6 17~ :!) 16 1,3
7 17~ :!) 48 6
8 17~ :!) 94 1,3
9 1&04 :!) 100 1,6
10 ]&04 :!) 60 1,6
11 1&04 :!) 60 1,3,4,5
12 1&04 :!) 100 1,3,4
13 (J7j(J) :!) 100 1,2,5,6
14 (J7j(J) :!) ]00 1,2,5,6
15 (J7j(J) :!) 20 1,2,3,4,5,6
16 (J7!OO :!) 20 1,3,4,6 Key to mangrove species: 1 = Bruguiera gymllorhiza; 2 = Lumllilzera liuorea; 3 :: Rhizophora samoensis; 4 = Rhizophora stylosa; 5 = Rhizophora x selala;
6 = Xylocarpus granatum.
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TABLE 3 Mangrove-associated algae at Ba and Rewa, and frequency of occurrence.
SITESIFREQUENCY Ba Rewa Mean % (F)
Number of pneumatophores sampled 320 320 RHODOPHYTA Bostrychia moritziafla (Sond. ex Klitz.) 15 31 28
J.Ag. Bostrychia radicans (Mont.) Mont. 41 29 35 Bostrychia simpliciuscula Harv. ex
lAg. J5 38 31 Bostrychia teflelLa (Lamx.) J .Ag. 22 11 Caloglossa sp. inedit (1)3 38 28 33 Caloglossa sp. inedit (2)4 15 25 20 Caloglossa leprieurii (Mont.) J.Ag. ~ 22 36 ErYlhrolrichia camea (Oillw.) J.Ag. :;n 29 24 Gelidium crillaLe(rurn.) Gaillon
var. perpusillum Pice. et Grunow' 38 17 27 Gelidium pusillum (Stackh.) Le Jol. 38 17 27 PolysiphOllia sp. 5 3 Rhodolachlle radicosa lton06 ~ 20 35 Stictosiphonia keLanensis (Grunow ex Post)
King et Puttock :D 29 30 PHAEOPHYfA Dictyola barlayresiafla Lamx. ~ 22 40 Dictyolopsis propagulifera W. Troll 15 10 13 CHLOROPHYfA ChLorodesmis hildebralldtii A.Gepp &
E.S. Gepp 18 59 38 Cladophoropsis sUlldallellsis Reinb .7 13 35 24
2Not previously recorded from Fiji. See Southet al., 1993. 3= CaLoglossa "adhaerens" sp. inedit. (King & Puttock, pers. comm.); probably previously recorded from Fiji asCalogLossa adllata (Zanard.) Howe (South & Kasahara, 1992). 4= Caloglossa "bengalensis" sp. inedit. (King & Puttock, pers. comm.). 'Not previously recorded from Fiji; see South et aI., (1993). 6'[0 be transferred to Bostrychia (Raj, 1993; Puttock, pers. comm.). 7Not previously recorded from Fiji ; see Southet aI., (1993) .
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TABLE 4
Summary of the presence (+) and absence (-) of algae according to host, Ba and Rewa, and number of epiphyte species per host.
HOST 1 2 3 4 5 67 8 9 10 11 12 13 14 15 16 17
BA B. gymflOrhiza + + - - + - + + + + + + + + + R.samoeflsis + + + - + + + + + + + + + + + + + R. stylosa + + + - + + + + + + + + + + + + + R. x selala + + + - + - + + + + + + + + X. granatum +
+ + - + - + + + + + + + +
REWA B.gymnorhiza + +++ + ++ + + + + + + + + + L. liuoralis + + + + + - + + + + + + + + + + R. samoeflsis + + + + + - + - + + + + + + + + R. stylosa + + + + + - + + + + + + + + + + R. xselala + + + + + + + + + + + + + + + + X. granatum + + + + + + + + + + + + + + + +
Key to species: 1 = Bostrychia moritziafla; 2 = B. radicalls; 3 = B. simpliciuscula; 4 = B. tene/fa; 5 = Calogfossa "adherefls"; 6 = C. "beflga/eflsis"; 7 = C. leprieurii; 8 = Erythrotrichia camea; 9 = GeLidiwn pusillum; 10 = G. criflale var. perpusillum; 11 = Polysiphonia sp.; 12 = Rhodolachfle radicosa (= Bostrychia sp.); 13 = Stictosiphoflia kelaneflSis; 14 = Dictyota bartayresiafla; 15 = Dictyotopsis propagulifera; 16 = Chlorodesmis hildebrafldtii; 17 = Cfadophoropsis sUlldalleflSis.
Total
12 16 16 12
12
16 15 14 15 16 16
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