Keppel et al.—Mesic of Dogotuki, Vanua Levu New Zealand Journal forest of Botany, 2006, Vol. 44: 273–292 0028–825X/06/4403–0273 © The Royal Society of New Zealand 2006 273 The flora, vegetation, and conservation value of mesic forest at Dogotuki, Vanua Levu, Fiji Islands GUNNAR KEPPEL Biology Division School of Biological, Chemical and Environmental Sciences Faculty of Science and Technology University of the South Pacific Suva, Fiji Present address: School of Integrative Biology, University of Queensland, St. Lucia, Brisbane, QLD 472, Australia. g.keppel@uq.edu.au ISAAC A. ROUNDS NUNIA T. THOMAS Institute of Applied Sciences University of the South Pacific Suva, Fiji Abstract The flora and vegetation of an area in north-eastern Vanua Levu that is part of Fiji’s last major system of mesic forest were studied and revealed a mosaic of vegetation types, including mesic sclerophyll forest, transition forest, stunted Dacrydium nidulum forest, mangrove forest, montane forest, brackish and freshwater wetlands, and disturbed landscapes. This is much more diverse than indicated by the previous “dry forest” label. The flora comprises more than 268 native species, several of which are rare or narrowly distributed. Relatively large and undisturbed stretches of vegetation and the presence of rare and novel species and vegetation types suggest that the landowners should be given every possible assistance in protecting this unique landscape. Keywords Dacrydium nidulum; Gymnostoma vitiense; Fagraea beteroana; Fiji; Melanesia; mesic forest; sclerophyll; South Pacific; transition forest; vascular flora; vegetation analysis B05035, Online publication date 13 September 2006 Received 16 August 2005; accepted 6 June 2006 INTRODUCTION Originating some 48 and 40 million years ago (Yan & Kroenke 1993), the Fiji Islands are an oceanic island group located between 15° and 22°S and 177°W and 177°E in the South Pacific Ocean (Fig. 1). The archipelago includes more than 500 islands and covers a territory of about 650 000 km2, less than 3% of which is land area. The two largest islands, Viti Levu (10 338 km2) and Vanua Levu (5535 km2) constitute more than 80% of the total landmass. On these two islands and some of the medium-sized islands, a pronounced windward-leeward effect is present due to orographic precipitation of moisture carried in the prevailing SE tradewinds, resulting in a wet SE side and a dry western side (Mueller-Dombois & Fosberg 1998). The vascular flora of Fiji has been well explored and described (Seemann 1865–73; Copeland 1929; Brownlie 1977; Smith 1979, 1981, 1985, 1988, 1991, 1996). In a review based on recent taxonomic literature, 137 families, 484 genera, and 1313 species of seed plants are recognised as indigenous to Fiji (Heads 2006). Although only 1 family and 11 genera are endemic to Fiji (Smith 1996), endemism at the species level has been estimated to be about 62% (Watkins 1995; Smith 1996). However, recent taxonomic revisions of various genera and families have reduced the number of species considered endemic to Fiji (Heads 2006). Mueller-Dombois & Fosberg (1998) recognised nine principal vegetation types in a synthesis based on several other treatments of Fiji’s vegetation (Smith 1951; Twyford et al. 1965; Cochrane 1969; Parham 1972; Berry & Howard 1973; Smith 1979; Ash 1992), including lowland rain forest, upland rain forest, cloud forest, dry forest, “talasiga” grasslands (a Fijian word meaning “sunburnt land”) freshwater wetland vegetation, mangrove forest and scrub, coastal strand vegetation, and smaller island vegetation. Tropical dry forest was first recognised by Twyford et al. (1965) as one of Fiji’s principal vegetation types in areas that receive 1750–2250 mm of rain. 274 New Zealand Journal of Botany, 2006, Vol. 44 Fig. 1 The Fiji Group. The rectangular area is enlarged in Fig. 2. Using aerial photographs and ground truthing, Berry & Howard (1973) recognised nine different forest types within this dry forest, most of which were dominated by Gymnostoma vitiense, Fagraea gracilipes, and Dacrydium nidulum (Appendix 1). Species of Myristica and Syzygium, Dysoxylum richii, Parinari insularum, and Agathis macrophylla may be locally dominant. In addition, a Sapindaceae pole forest and a woodland dominated by introduced raintrees (mostly Samanea saman and Albizia lebbeck) was recognised. Mueller-Dombois & Fosberg (1998) suggested that this dry forest should be classified as a mesic forest on a global scale. Recent discovery of a “true” dry forest, which has several deciduous species, supports this. Therefore, we will henceforth refer to the dry forest of Twyford et al. (1965), Berry & Howard (1973), and Mueller-Dombois & Fosberg (1998) as “mesic forest” (MF). MF has mostly been replaced by talasiga grasslands (Smith 1951; Parham 1972) and is now restricted to small areas on Vanua Levu (Berry & Howard 1973). The need to protect the remaining MF was recognised when a proposed system of national parks was developed for the Fiji Islands (Lees 1989). In particular, the MF around and between the Vunivia and Nasavu Catchments, Fiji’s easternmost river systems, on the lower Udu Peninsula (Vanua Levu), about 45 km east-north-east of Labasa (Fig. 1, 2), was identified as a possible heritage site and national park. Lees (1989) did not provide a species list but noted the presence of well-developed mangrove forest. D’Espessis (1941 in Bogiva 1994) was the first to comment on the unique vegetation and physical and climatic features of the Udu Peninsula. Subsequent evaluations of the area all strongly supported the establishment of a national park in the Vunivia and Nasavu Catchments (Weaver 1992a,b; Bogiva 1994). A preliminary survey of the existing archaeological and historical literature also identified the area as potentially interesting (Chabaniuk 1992). Despite the seeming uniqueness of the area, no detailed Keppel et al.—Mesic forest of Dogotuki, Vanua Levu 275 Fig. 2 Nasavu and Vunivia Catchment systems and locations of the study sites. scientific survey of the flora and fauna of either the catchments or the MF has been conducted; neither is any MF in Fiji protected, although the need for protection was highlighted by Lees (1989). We were approached by members of one of the landowning clans (Kawa nei Tagi Tuba) to substantiate the assertion that the two catchments are botanically unique, to allow them to consider conservation measures. This paper presents the results of this study and, therefore, constitutes the first detailed description of the structure, composition, and diversity of MF in Fiji. It is also the first extensive survey of the lower Nasavu Catchment and findings are related to the conservation importance of the proposed Vunivia and Nasavu catchment forest reserve. It is hoped that this study is a step towards a complete survey of the flora and fauna in the two catchment areas and their eventual protection. METHODS Study area The area studied is the Dogotuki District, Macuata, which is part of the proposed Vunivia and Nasavu Catchment Reserve and is on the lower reaches of the Nasavu and Vunivia Rivers (Fig. 2). At the nearby Udu Point weather station little seasonality and a total annual precipitation of 2450 mm are recorded (Fig. 3). However, the precipitation regime at the study site is likely to differ, because much of the moisture carried by the SE tradewinds is likely to precipitate over the peninsula, thereby creating a minor rainshadow on the northward side of the peninsula. We expect the actual climate at the study side to be similar to that for Udu Point but to have a lower rainfall curve, resulting in a lower total annual precipitation. New Zealand Journal of Botany, 2006, Vol. 44 276 Flora Research was carried out in July 2003. If possible, we identified species in the field using Smith (1979, 1981, 1985, 1988, 1991, 1996), Brownlie (1977), Whistler (1992, 1995, 2000), Purseglove (1972, 1974), and Keppel & Ghazanfar (in press). We also collected other species for identification and deposition at the South Pacific Regional Herbarium (SUVA), Suva, Fiji. Vegetation A reconnaissance by walking through the area identified 10 putatively unique vegetation types (Mueller-Dombois & Ellenberg 2002) (Table 1): 5 types of undisturbed, terra firma forests (“Bainivara”, “Bereniyalo”, “Caudramea”, “Nabourewa”, “Nautuutu”), disturbed terra firma forest (“Dareka”), mangrove forest (“mangroves”), fire-disturbed savanna (“Bainivara Dist”), and two herbaceous swamp communities (“mangroves”, “Waitabua”). In each putatively unique vegetation type a species list was prepared, except for the mangrove back swamp, which could not be properly surveyed because of the swampy conditions and was therefore excluded from the analyses. At each site with undisturbed terra firma forest, we set up a 10 × 60 m plot in a representative, homogenous portion to document the structure and composition of the vegetation present. In each plot we recorded the dbh, bole height, crown height, crown width, and position of every tree with dbh ≥ 10 cm. We also recorded climbers, epiphytes, and species in the understorey. Table 1 Analysis We assessed the uniqueness of the different putative vegetation types using agglomerative clustering of the Community Analysis Package (CAP; Henderson & Seaby 2002) using average linkage based on the Jaccard coefficient. CAP was also used for a twoway indicator species analysis (TWINSPAN) to identify the species characteristic for each vegetation type. Data sets used for analysis are available from GK upon request. In addition, profile diagrams for the five sites with 10 × 60 m plots were drawn. RESULTS Flora A total of 336 species were recorded, 268 (80%) of which are indigenous (Table 2; Appendix 2). This list is comprehensive but by no means complete. Endemism of the native flora is about 34% (90 species). Rubiaceae is the most diverse family, with 20 native species, followed by Euphorbiaceae and Orchidaceae with 13 species each and Cyperaceae with 10 species. Other common families include Moraceae, Myrtaceae, Poaceae, and Sapotaceae (Table 3; Appendix 2). Several of the endemic species are rare or have a narrow distribution. The endemic grass Leptaspis angustifolia has a very narrow range, restricted to the Macuata Province on Vanua Levu. Plants that are only found on Vanua Levu (and in some cases also on the adjacent island of Taveuni) are: Location and vegetation of study sites. See Fig. 2 for location of locality names. Site Location Bainivara Bainivara Dist Bainivara Bainivara Bereniyalo (“footprints of the spirit”) Caudramea Dareka Bereniyalo Caudramea Dareka Mangrove Forest Mangrove Back Swamp Nabourewa Nautuutu Waitabua Dareka Dareka Nabourewa Nautuutu Waitabua Vegetation of plot mesic forest (c. 15° slope) savanna with remnant trees of mesic forest and fire resistant species, several trunks charred, indicating recent fire. stunted forest mesic forest (ridge) disturbed mesic forest adjacent to Dareka settlement and Dogotuki village mangroves herbaceous swamp forest with elements of mesic and rain forest forest with elements of mesic and rain forest lake with mainly herbaceous swamp vegetation on margins Keppel et al.—Mesic forest of Dogotuki, Vanua Levu 277 Fig. 3 Climate diagram of the weather station at Udu Point from 1994 to 1999 (Fig. 2). Altitude: 62 m. Average annual temperature: 26°C. Average annual rainfall: 2451 mm. Table 3 List of all families with more than five indigenous species identified, stating the number of native and introduced species. Family Indigenous* Introduced Ferns and Fern Allies Gymnosperms Dicotyledons Monocotyledons TOTAL Endemic Table 2 Summary of the classification, origin and distibution of the species identified. *, includes endemic species. 4 42 - 42 78 8 90 4 177 45 268 51 17 68 4 228 62 336 TOTAL Rubiaceae Euphorbiaceae Orchidaceae Cyperaceae Poaceae Moraceae Myrtaceae Sapotaceae Apocynaceae Caesalpiniaceae Meliaceae Rhamnaceae Rhizophoraceae Verbenaceae Adiantaceae Anacardiaceae Clusiaceae Davalliaceae Indigenous Introduced Total 20 13 13 10 8 8 8 7 6 6 6 6 6 5 5 5 5 5 4 5 – 2 7 1 1 – 4 1 – – – 2 – – – – 24 18 13 12 15 9 9 7 10 7 6 6 6 7 5 5 5 5 278 New Zealand Journal of Botany, 2006, Vol. 44 Fig. 4 Grouping of the nine sample sites using agglomerative clustering and average linkage based on Jaccard coefficient. Cyathocalyx vitiensis (Annonaceae), Parsonsia smithii (Apocynaceae), Veitchia filifera (Arecacaceae), Macaranga membranacea (Euphorbiaceae), Airosperma vanuense, Ixora myrtifolia, and Psychotria argantha (all Rubiaceae). Three additional species were classified as rare because there are 5 or fewer cited specimens in Smith’s (1979, 1981, 1985, 1988, 1991, 1996) Flora Vitiensis Nova: Garnotia linearis (Poaceae, 5 collections), Maniltoa vestita (Caesalpiniaceae, 3 collections), and Syzygium simillimum (Myrtaceae, 2 collections). In addition, one of the Psychotria (Rubiaceae) collected from Mt Cavanavula, differs from all other species in the genus described from Fiji and is possibly new to science. Vegetation Jaccard coefficients ranged between 0 (no similarity) and 0.25 (Fig. 4), showing low similarities between the different putative vegetation types, confirming our assessment based on visual reconnaissance and entitation. TWINSPAN produced a dendogram with clusters similar to those of the cluster analysis in Fig. 4 and is not shown. However, we used the TWINSPAN output to determine characteristic species for the various clusters. For example, wetlands are clearly differentiated from forests, sharing no common species. The wetland species Acrosticum aureum, Stenochlaena palustris, and Pandanus tectorius were characteristic for the two wetlands (“mangroves” and “Waitabua”), which had low similarities (J = 0.2). The fire-disturbed vegetation at “Bainivara Dist” and the stunted vegetation of “Bereniyalo” are distinct (Fig. 4). Several species that in Fiji are often associated with frequent fires, such as Casurina equisetifolia, Dodonea viscosa, Pennisetum polystachyon, and Pteris ensiformis, are only recorded from “Bainivara Dist”. Early successional species, such as Alstonia pacifica, Commersonia bartramia, and Geniostoma rupestre were unique to “Dareka”, which is possibly related to recent swidden agriculture, e.g., we found a small plantation of kava (Piper methysticum) in one location. Other species unique to “Dareka” are Centotheca lappacea, Dichapetalum vitiense, Exocarpus vitiensis, Serianthes melanesica, and Adenanthera pavonia. Because our study focused on native, undisturbed vegetation types, we considered the vegetation on “Bainivara Dist” and “Dareka” together as “disturbed vegetation”. “Bereniyalo” has stunted vegetation characterised by several unique species, including Acacia richii, Gahnia aspera, Garcinia adiantha, Hibbertia luccens, and Leucopogon septentrionalis, and is considered a separate vegetation type. In addition, the plots of “Bainivara” and “Caudramea” form a distinct cluster and are here considered as mesic forest (MF). They uniquely share Alpinia vitiensis, Syzygium decussatum, Connarus pickeringii, Phyllanthus heterodoxus, Podocarpus neriifolius, Syzygium fijiense, Tapeinospermum grande, Dacrydium nidulum, and Fagraea beteroana. “Nautuutu” and “Nabourewa” also form a distinct cluster and are here considered to be “transition forest” (TF) because they have physiognomic and taxonomic characteristics that are intermediate between MF and lowland tropical rain forest (see below). Only Buchanania vitiensis, Cordyline fruticosa, Dillenia biflora, and Garcinia pseudoguttifera are unique to this group, showing that it is less clearly defined. Gymnostoma vitiense and the climbers Freycinetia impavida, Flagellaria gigantea, Flagellaria indica, and Lygodium reticulatum commonly occur in both MF and TF. An additional vegetation type, montane forest, is recognised on Mt Cavanavula based on specimens collected and vegetation descriptions. However, this vegetation type was not studied in detail. Therefore, we identified eight vegetation types. 1. Mesic forest (MF) This is the most common vegetation type in the study area and is dominated by Gymnostoma vitiense, Dacrydium nidulum, and Fagraea gracilepes, which are present in different proportions in different locations. The forest is similar to but more diverse (Fig. 5) than the “dry forest” of Berry & Howard (1973). Species composition differs on ridges (Fig. 6), with gymnosperms being more diverse and dominant and the legume Intsia bijuga becoming an important component of the forest. This forest type lacks deciduous species but about a quarter of its species are sclerophyllous. 279 Fig. 5 Mesic slope forest. Profile diagram of trees with dbh ≥10 cm in a 60 × 10 m plot on a c. 15° slope at Bainivara, Dogotuki, Macuata, Vanua Levu. Buat, Buchanania attenuata; Cavi, Canarium vitiense; Crha, Crossostylis harveyi; Cyin, Cyathocalyx insularis; Gmvi, Gmelina vitiensis; Gyvi, Gymnostoma vitiense; Hafl, Haplolobus floribundus; Magr, Maniltoa grandiflora; Mygi, Myristica gillespieana; Pafi, Palaquium fidjiense; Poga, Pouteria garberi; Plti, Pleiogynium timoriense; Syfi, Syzygium fijiense; Sysp, Syzygium sp. Keppel et al.—Mesic forest of Dogotuki, Vanua Levu 280 New Zealand Journal of Botany, 2006, Vol. 44 Fig. 6 Mesic ridge forest. Profile diagram of trees with a dbh of 10 cm or more in a 30 × 10 m plot on a ridge at Caudramea, Dogotuki, Macuata, Vanua Levu. Agma, Agathis macrophylla; Cace, Calophyllum cerasiferum; Dani, Dacrydium nidulum; Dysp, Dysoxylum sp.; Gyvi, Gymnostoma vitiense; Inbi, Intsia bijuga; Madi, Manilkara dissecta; Syfi,Syzygium fijiense; Syru, Syzygium rubescens; Sysp, Syzygium sp. 2. Stunted Dacrydium nidulum forest This is an interesting variant of MF and is found at “Bereniyalo”, where Dacrydium nidulum forms low-growing, monodominant stands (Fig. 7). In some places only a few sedges or no vegetation at all persist. Gahnia aspera, Garcinia adiantha, Garcinia myrtifolia, Hibbertia luccens, Leucopogon sepentrionalis, and Lycopodiella cernua are restricted to this vegetation type. associated species (Acrostichum aureum, Dalbergia candatensis, Excoecaria agallocha, Lumnitzera littorea, species of Xylocarpus). In addition to these typical mangrove species, the climbers/epiphytes Abrus precatorius, Davallia solida, Derris trifoliata, and Hoya australis were restricted to mangrove vegetation. The mangrove forest is extensive, intact, and unique because trees of the lowland forest, such as Agathis macrophylla, are found on the inner margins of this forest in some places. 3. Transition forest (TF) This has a species composition intermediate between MF and the lowland rainforest described by Mueller-Dombois & Fosberg (1998). It contains several species (Amaroria soulameoides, Dillenia biflora, Garcinia pseudoguttifera, Gironniera celtidifolia) that are usually associated with lowland rainforest and species (Fagraea gracilipes, Gymnostoma vitiensis) that are dominant in MF. In addition this forest has some common rainforest features (stilt roots, buttresses, palms; Fig. 8, 9). The composition of transition forest seemingly varies greatly. For example, while the forest at “Nabourewa” is dominated by Gymnostoma vitiensis (Fig. 8), that at “Nautuutu” is dominated by Myristica gillespieana (Fig. 9). 5. Brackish swamps These are mainly herbaceous communities found on the inner margins of mangroves and dominated by Eleocharis dulcis with occasional trees of Pandanus tectorius and, rarely, Barringtonia racemosa. The inner margins of these swamps, which presumably have a very low salinity, have been used to plant dalo (Colocasia esculenta; also known as taro), a staple food. Such areas differ in species composition, with native (Acrostichum aureum, Diplazium esculentum, Nephrolepis biserrata, Barringtonia racemosa) and invasive (Ludwigia occidentalis, Mikania micrantha, Paspalum conjugatum, Merremia peltata) species codominating. 4. Mangrove forest This covers almost the entire coastline. It is composed of coastal Rhizophora (R. stylosa, R. mangal, and the sterile hybrid R. ×selala) formations and landward Brugiera gymnorrhiza formations and 6. Freshwater wetlands A freshwater lake, Waitabua, was the only freshwater wetland in our study area. Its centre is open water, which is surrounded by a band of Eleocharis dulcis (Cyperaceae) up to 5 m wide. On the periphery of this lake is a very thin band (<50 cm wide) consisting of several other sedges, the ferns Acrostichum aureum and Stenochlaena palustris, and Pandanus tectorius. Several of the epiphytic orchids were collected from trees facing this inland lake, and Eleocharis dulcis and Rynchospora corymbulosa are unique to this vegetation. 7. Montane forest This is restricted to the top of Mt Cavanavula (c. 400 m), as indicated by the collection of species typical of montane vegetation. In addition the forest is stunted and gnarled (M. Fox, D. Jackson, and L. Cokanasiga pers. comm.), which is typical of montane or cloud forest. The mountain is isolated and this suggests that it could have new, endemic species, an assertion substantiated by the collection of a species of Psychotria aff. macrocalyx (Rubiaceae) not listed in Smith (1988) and different from Psychotria volii (Gardner 1997). 8. Disturbed vegetation and anthropogenic landscapes These have been caused by five major factors (fire, agriculture, settlements, cyclones, logging). While agriculture and settlements are more permanent disturbances that result in complete transformation of the original vegetation, other disturbances are usually followed by gradual recovery. However, food gardens often replace logged forests. Agriculture may have several effects on the vegetation, depending on the duration and extensiveness of cultivation. Swidden agriculture at “Dareka” appeared to maintain forest cover but to change species composition. Extensive agriculture over long periods of time was seen to result in soil degradation and dominance of introduced species. A few uprooted trees and several broken branches, likely the result of Cyclone Ami which passed close by the study site in January 2003, covered the forest floor in some locations. Some locations showed evidence of relatively recent fires (e.g., “Bainivara Dist”), and a displacement of MF species by more fire-resistant species such as Casurina nodiflora and Dodonaea viscosa was observed (both of which were unique to the “Bainivara Dist” side in the TWINSPAN analysis). Bracken ferns (Dicranopteris linearis, Gleichenia oceanica, and species of Pteris) dominate this landscape. 281 Fig. 7 Stunted Dacrydium nidulum forest. Profile diagram of trees with dbh ≥10 cm in a 60 × 10 m plot of stunted Dacrydium nidulum forest on a flat at Bereniyalo, Dogotuki, Macuata, Vanua Levu. Acri, Acacia richii; Dani, Dacrydium nidulum; Gaad, Garcinia adiantha; Gyvi, Gymnostoma vitiense; Madi, Manilkara dissecta; Pone, Podocarpus nerifolius; Sysp, Syzygium sp. Keppel et al.—Mesic forest of Dogotuki, Vanua Levu 282 New Zealand Journal of Botany, 2006, Vol. 44 Fig. 8 Transition forest. Profile diagram of trees with dbh ≥10 cm in a 60 × 10 m plot on a c. 25° slope at Nabourewa, Dogotuki, Macuata, Vanua Levu. Buvi, Buchanania vitiensis; Crha, Crossostylis harveyi; Dibi, Dillenia biflora; Fagr, Fagraea gracilipes; Gasp, Gardenia sp.; Gyvi, Gymnostoma vitiense; Myca, Myristica castaneifolia; Mygi, Myristica gillespieana; Pain, Parinari insularum; Syru, Syzygium rubescens; Vefi, Veitchia filifera. Keppel et al.—Mesic forest of Dogotuki, Vanua Levu 283 Fig. 9 Transition forest. Profile diagram of trees with dbh ≥10 cm in a 50 × 10 m plot on a c. 10° slope at Nautuutu, Qaranivai, Macuata, Vanua Levu. Amso, Amaroria soulameoides; Anlu, Anacolosa lutea; Buvi, Buchanania vitiensis; Dibi, Dillenia biflora; Gaps, Garcinia pseudoguttifera; Inbi, Intsia bijuga; Mygi, Myristica gillespieana; Pafi, Palaquium fidjiense; Sysp, Syzygium sp.; Vaam, Vavaea cf. amicorum; Vefi, Veitchia filifera. 284 DISCUSSION The flora of Dogotuki, with 274 native species (about 34% are endemic), is diverse. Tuiwawa (1999) found a more diverse flora (366 native species), similar percentage of indigenous species (86%), but much higher percentage endemism (49%) in moist rainforest on Viti Levu. This may indicate that the endemism in Fiji’s MF is lower than in the wet forests and corresponds to the theory of increased diversity with increased precipitation (Gentry 1988). The flora includes 12 rare or narrowly distributed species. In addition, some species, such as Hibbertia lucens and Sacromelicope petiolaris, are mainly restricted to dry zone habitats and are, therefore, likely to be endangered through habitat destruction. MF is more diverse than anticipated based on previous work. For example, the profile diagram of MF by Barry & Howard (1973; reproduced in Mueller-Dombois & Fosberg 1998, p. 124) has only five labelled taxa and Fagraea gracilepes is much more common than in our plots. Relationships between the different forest sites portrayed by the dendogram based on agglomerative clustering (Fig. 4) and TWINSPAN (similar to Fig. 4 but not shown) must be considered with caution, because they are solely based on presence/absence data and do not take into account the abundance of the different species. In particular, “Bereniyalo”, which is almost monodominant, and “Dareka”, which has a great number of unique species, may have clustered differently if sufficient abundance data had been collected. The previously reported homogenous MF (Lees 1989) of the Vunivia and Nasavu catchments is here shown to be a unique and diverse mosaic of different vegetation types, at least in the lower Nasavu catchment. This mosaic is likely to be caused by a combination of topographical, microclimatic, and fine-scale geological differences. Differences in species composition were so pronounced that we divided MF into three distinct forest types: MF, TF, and stunted Dacrydium nidulum forest. While MF is the predominant forest type, other vegetation types, the extent of which should be mapped, cover substantial areas. The heterogeneity of Fiji’s MF was first recognised by Berry & Howard (1973). They reported eight different forest types, most of them dominated by Gymnostoma vitiense, Dacrydium nidulum, Fagraea gracilepes, or a combination of these species (Appendix 1). These three species or a combination of two of them also dominate the MF in this study. New Zealand Journal of Botany, 2006, Vol. 44 In some places MF has been removed for logging and agricultural purposes or disturbed by burning. There are also pockets of an apparently moister forest type (TF), which have species common in tropical lowland rainforest in addition to those characteristic of MF. An interesting feature is the abundance of conifers, mainly Dacrydium nidulum and Agathis macrophylla, at low altitudes. Monodominant stands, such as the stunted Dacrydium nidulum forest at Bereniyalo, are rare in the tropics and can be caused by nutrient-poor soils or compounds impairing plant growth (Richards 1996). In the Pacific, monodominant forests have been found in mangroves (Thaman et al. 2005), swamps (Whitmore 1969; Ash & Ash 1984), and some terra firma sites, some of which, but not all, have a history of previous anthropogenic disturbance (Whitmore 1969; Bayliss-Smith et al. 2003). Further studies are required to determine the causative factors of this stunted, monodominant growth. In addition to these vegetation types, there are large stretches of mangrove forest along the coast and several wetlands. The latter includes brackish swamps bordering mangroves and inland lakes. Wetlands are generally highly disturbed in Fiji (Ash & Ash 1984) and in Oceania (Scott 1993). Mangrove forests have disappeared rapidly in Fiji over recent years and are now considered to require management (Lal 1990). We found extensive and relatively intact mangrove forests (similar in structure and composition to those described by Ghazanfar et al. (2001) and Thaman et al. (2005)) that, like the other wetlands, had few invasive species. In some places, trees usually associated with terra firma forest grew on the inner margins of mangrove forests, a phenomenon previously observed by IAR in some places along the SE coast of Viti Levu. MF was probably very common on Viti Levu and Vanua Levu in prehistoric times (Southern 1986) but has been reduced to small remnant pockets on Viti Levu and Vanua Levu and is possibly entirely extirpated on the former island (Twyford et al. 1965; Berry & Howard 1973). The MF in the Vunivia and Nasavu Catchments is the last extensive stand of this vegetation type remaining in Fiji (Lees 1989). Extensive, well-developed stretches of mangrove forest are also found. In addition, we observed herbaceous freshwater and brackish wetlands that had very few invasive species and a stunted forest dominated by Dacrydium nidulum. The latter forest type has not been previously reported. We also identified several species that are rare or have a restricted distribution. All these findings underline the conservation Keppel et al.—Mesic forest of Dogotuki, Vanua Levu value of the area. We therefore concur with previous authors (d’Espessis 1941 in Bogiva 1994; Lees 1989; Weaver 1992a,b; Bogiva 1994) regarding the uniqueness of the vegetation of the Vunivia and Nasavu Catchments. At present, logging is the single largest threat to this forest system. Several forests in close vicinity have already been cut and there is considerable interest in continuation of timber extraction (E. Kosoluva & Y. Roberts pers. comm.). While logging would provide cash for the local communities, it would also damage the forest almost irreversibly. Although no data on regeneration of mesic forest are available, it is likely, based on the slower growth rates of forests in drier regions, that it would take longer than for lowland tropical rainforest, which requires several centuries (Riswan et al. 1985). Other threats are invasive plant species, with the raintrees (Albizia lebbeck and Samanea saman) and the African tulip tree (Spathodea campanulata) posing the biggest threats. Recent work by Rolett & Diamond (2004) has pointed towards a high susceptibility of dry zone vegetation to disturbance. Thus, the demise of MF in Fiji probably commenced soon after the arrival of human beings. This study shows that botanical surveys can be a powerful tool in conservation and environmental management. By identifying species and vegetation types, richness in species and vegetation can be determined, species that are rare or have a restricted range identified, and spots with unique vegetation located. This is especially true on isolated oceanic islands like Fiji, where the fauna is depauperate (Flannery 1995; Watling 2001; Morrison 2003) and the flora relatively well known. Considering the uniqueness and the relatively pristine condition of the Nasavu and Vunivia catchment, the protection of its ecosystems and biodiversity should be considered a conservation priority. Although the site has been proposed to become a forest reserve (Lees 1989) and was, after work by Weaver (1992b), incorporated into a list of priority areas for protection under Fiji’s National Environment Strategy (Watling & Chape 1993), the proposed reserve is still not formalised and logging is occuring within its intended boundaries. As already mentioned by Weaver (1992b), protection would require careful consideration of the needs of landowners. A possible way of addressing this would be by creating a mosaic of protected (pristine) and usage (disturbed) areas. In addition, alternative sources of income for the inhabitants of the two catchments need to be ensured. Any road 285 construction or development projects should be carefully planned to avoid the introduction and spread of alien species. ACKNOWLEDGMENTS This study was made possible by Emma Adi Kosoluva and Yvonne Roberts, who hosted the visit and covered the travel expenses of the team members. 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Forest type Sapindaceae pole forest Buabua forest Moderately stocked velau-yaka dry-zone forest Low-stocked dryzone forest on long, very steep slopes Location Dominant species VanL, Kan Gymnostoma vitiense, Aryterya brackenridgei, Vavaea amicorum, Elattostachys falcata, Dysoxylum spp., Endospermum spp. VanL Fagraea gracilipes, Myristica spp., Syzygium spp. VanL Gymnostoma vitiense, Dacrydium nidulum, Parinari insularum, Calophyllum spp., Syzygium spp. VanL Gymnostoma vitiense, Calophyllum spp., Syzygium spp., Agathis macrophylla Kaudamu-velau woodland VanL, ViL Myristica spp., Gymnostoma vitiense, (lim) Syzygium spp., Intsia bijuga, Parinari insularum, Agathis macrophylla Raintree woodland Velau woodland Moderately stocked velau-buabua forest VanL, ViL Albizia lebbeck, Samanea saman VanL, ViL Gymnostoma vitiense, Acacia richii, Dacrydium nidulum, Myristica spp., Fagraea gracilipes, Dysoxylum richii, Endospermum spp., Garcinia pseudoguttifera, Semecarpus vitiensis VanL Gymnostoma vitiense, Fagraea gracilipes, Dysoxylum richii Velau-vesi forest VanL Gymnostoma vitiense, Intsia bijuga, Parinari insularum, Canarium spp. Other common species and genera Palaquium spp., Parinari insularum, Dysoxylum richii Canarium spp., Parinari insularum, Palaquium porphyreum, Palaquium spp., Myristica spp., Serianthes melanesica, Intsia bijuga, Dacrydium nidulum, Fagraea gracilipes Canarium spp., Palaquium porphyreum, Palaquium spp., Calophyllum spp., Dysoxylum richii, Endospermum spp., Xylopia pacifica Parinari insularum, Canarium spp., Palaquium spp., Agathis macrophylla, Podocarpus neriifolius, Brackenridgea nitida, Crossostylis pedunculata, Sapindaceae Parinari insularum, Syzygium spp., Dacrydium nidulum, Podocarpus neriifolius, Intsia bijuga, Trichospermum spp. New Zealand Journal of Botany, 2006, Vol. 44 288 Appendix 2 Annotated checklist of vascular plants. All species are indigenous, unless marked. E, endemic to Fiji; *, recently (historically) introduced species LOMARIOPSIDACEAE Lomagramma polyphylla Brack. PTERIDOPHYTA (Ferns and Fern Allies) LYCOPSIDA LYCOPODIACEAE Lycopodinella cernuum (L.) Pic.Serm. Huperzia phlegmaria (L.) Rothm. Huperzia squarrosa (G.Forst.) Trev. MARATTIACEAE Angiopteris evecta (Forst.) Hoffm. SELAGINELLACEAE Selaginella breynioides Baker E Selaginella laxa Spring Selaginella viridangula Spring E FILICOPSIDA ADIANTACEAE Taenitis pinnata (J.Sm.) Holttum var. pinnata POLYPODIACEAE Drynaria rigidula (Sw.) Bedd. Pyrrosia lanceolata (L.) Farw. Microsorum grossum (Langsd. & Fisch.) S.B.Andrews PTERIDACEAE Acrostichum aureum L. Pteris ensiformis Burm. Pteris pacifica Hieron. SCHIZAEACEAE Lygodium reticulatum Schkuhr Schizaea dichotoma (L.) J.E.Sm. ASPIDIACEAE Tectaria latifolia (G.Forst.) Copel. THELYPTERIDACEAE Sphaerostephanos invisus (G.Forst.) Holtt. ASPLENIACEAE Asplenium australasicum Hook. Asplenium polydon G.Forst. VITTARIACEAE Halopteris elongata (Sw.) H.E.Crane Monogramma acrocarpa (Holttum) D.L.Jones ATHYRIACEAE Diplazium esculentum (Retz.) Sw. Callipteris prolifera (Lam.) Bory GYMNOSPERMAE (Gymnosperms) CONIFERALES ARAUCARIACEAE Agathis macrophylla (Lindl.) Masters BLECHNACEAE Blechnum orientale L. Stenochlaena palustris (Burm.) Bedd. CULCITACEAE Calochlaena straminea (Labill.) M.D.Turner & R.A.White CYATHACEAE Cyathea lunalata Copel. Cyathea propinqua Mett. E DAVALLIACEAE Davallia solida var. fejeensis (Hook.) Nootenboom Davallia solida (G.Forst.) Sw. var. solida Humata heterophylla (Sm.) Desv. Nephrolepis biserrata (Sw.) Schott Nephrolepis hirsutula (G.Forst.) C.Presl. DENNSTAEDTIACEAE Pteridium esculentum (Forst.) Cockayne GLEICHENIACEAE Dicranopteris linearis (Burm.f.) Underw. Gleichenia oceanica Kuhn HYMENOPHYLLACEAE Cephalomanes atrovirens (Kunze) Copel. Selenodesmium dentatum (Bosch) Copel. Nesopteris intermedia (Bosch) Copel. LINDSACEAE Tapeinidium melanesicum Kramer PODOCARPACEAE Dacrydium nidulum de Laub. Podocarpus neriifolius D.Don GNETALES GNETACEAE Gnetum gnemon L. ANGIOSPERMAE (Angiosperms) DICOTYLEDONAE (Dicotyledons) ACANTHACEAE Graptophyllum insularum (A.Gray) A.C.Sm. Pseuderanthemum laxiflorum (A.Gray) C.E.Hubb. E *Hemigraphis alternata T.Anders. AMARANTHACEAE *Alternanthera sessilis (L.) R.Br. ex Roem. & Schult. *Celosia argentea L. ANACARDIACEAE Buchanania attenuata A.C.Sm. E Buchanania vitiensis Engl. E *Mangifera indica L. Pleiogynium timoriense (DC.) Leenh. Semecarpus vitiense (A.Gray) Engl. ANNONACEAE Cyathocalyx cf. vitiensis A.C.Sm. E APOCYNACEAE *Allamanda cathartica L. Alstonia pacifica (Seem.) A.C.Sm. Keppel et al.—Mesic forest of Dogotuki, Vanua Levu Alstonia costata (G.Forst) R.Br. Alyxia stellata (J.R. & G.Forst.) Roem. & Schult. *Catharanthus roseus ( L.) G.Don Cerbera manghas L. Ervatamia obtusiuscula Markgraf Parsonsia smithii (A.Gray) Markgraf E *Plumeria obtusa L. *Plumeria rubra L. ARALIACEAE Plerandra vitiensis (Seem.) Baill. E Polyscia multijuga (A.Gray) Harms *Polyscias scutellaria (Burm.f.) Fosberg ASCLEPIADACEAE Hoya australis R.Br. Tylophora sp. ASTERACEAE *Bidens pilosa L. *Conyza bonariensis (L.) Cronquist *Mikania micrantha H.B.K. *Pseudoelephantopus spicatus (Juss. ex Aubl.) C.F.Baker *Synedrella nodiflora (L.) Gaertn. *Vernonia cinerea (L.) Less. BIGNONIACEAE *Spathodea campanulata Beauv. BORAGINACEAE Cordia subcordata Lam. BURSERACEAE Canarium cf. vitiense A.Gray Haplolobus floribundus subsp. salomonensis (C.T.White) Leenh. CAESALPINIACEAE Caesalpinia bonduc (L.) Roxb. Cynometra insularis A.C.Sm. E Intsia bijuga (Colebr.) Kuntze Kingiodendron platycarpum Burtt E Maniltoa grandiflora (A.Gray) Scheffer Maniltoa vestita A.C.Sm. E *Senna alata (L.) Roxb. CARICACEAE *Carica papaya L. CASUARINACEAE Casuarina equisetifolia L. Gymnostoma vitiense L.A.S.Johnson E CHRYSOBALANACEAE Atuna racemosa Raf. Parinari insularum A.Gray CLUSIACEAE Calophyllum cerasiferum Vesque E Calophyllum vitiense Turr. E Garcinia adiantha A.C.Sm. & S.Darwin E Garcinia myrtifolia (A.Gray) Seem. Garcinia pseudoguttifera Seem. 289 COMBRETACEAE Lumnitzera littorea (Jack) Voigt Terminalia catappa L. CONNARACEAE Connarus pickeringii A.Gray E CONVOLVULACEAE Ipomoea indica (Burm.) Merr. Ipomoea littoralis Bl. Merremia peltata (L.) Merr. CUNNONIACEAE Geissois ternata A.Gray E DICHAPETALACEAE Dichapetalum vitiense (Seem.) Engl. DILLENIACEAE Dillenia biflora Martelli Hibbertia luccens Brogn. & Gris ex Sébert & Pancher ELAEOCARPACEAE Elaeocarpus storckii Seem. E EUPHORBIACEAE Acalypha repanda var. denudata (Muell.Arg.) A.C.Sm. *Acalypha wilkesiana Roxb. f. wilkesiana Baccaurea stylaris Muell.Arg. E *Chamaesyce hirta (L.) Millsp. Claoxylon echinospermum Muell.Arg in DC. E Claoxylon fallax Muell.Arg. *Codiaeum variegatum (L.) Jussieu Endospermum macrophyllum (Muell.Arg.) Pax & Hoffm. E Excoecaria agallocha L. Glochidion amentuligerum (Muell.Arg) Croizat E Glochidion cordatum Seem. E Homalanthus nutans Benth. & Hook.f. ex Drake Macaranga membranacea Muell.Arg. E Macaranga seemannii var. capillata A.C.Sm. Macaranga vitiensis Pax & Hoffm. E *Manihot esculenta Crantz Phyllanthus heterodoxus Muell.Arg. E *Phyllanthus virgatus G.Forst. FABACEAE Dalbergia candenatensis (Dennst.) Prain *Derris malaccensis (Benth.) Prain Derris trifoliata Lour. *Desmodium triflorum (L.) DC. Inocarpus fagifer (Parkinson) Fosberg *Pueraria lobata (Willd.) Ohwi Vigna marina (Burm.) Merr. FLACOURTIACEAE Flacourtia subintegra A.C.Sm. E Homalium vitiense Benth. E LAMIACEAE *Hyptis pectinata. (L.) Poit. *Ocimum basilicum L. LAURACEAE Cassytha filiformis L. 290 New Zealand Journal of Botany, 2006, Vol. 44 LECYTHIDACEAE Barringtonia edulis Seem. Barringtonia racemosa (L.) Spreng. Maesa insularis Gillespie E Maesa persicifolia A.Gray E Tapeinosperma grande (Seem.) Mez E LOGANIACEAE Fagraea berteroana A.Gray ex Benth. Fagraea gracilipes A.Gray Geniostoma rupestre J.R. & G.Forst. Neuburgia cf. corynocarpa (A.Gray) Leenh. MYRTACEAE Decaspermum vitiense (A.Gray) Niedenzu E *Psidium guajava L. Syzygium curvistylum (Gillespie) Merr. & L.M.Perry Syzygium decussatum (A.C.Sm.) Biffin & Craven E Syzygium effusum (A.Gray) C.Muell. Syzygium eugenoides (Merr. & L.M.Perry) Biffin & Craven E Syzygium fijiense L.M.Perry E Syzygium rubescens (A.Gray) C.Muell. E Syzygium simillimum Merr. & L.M.Perry E LORANTHACEAE Decaisnina forsteriana (Schult. & Schult.f.) Barlow LYTHRACEAE *Cuphea carthagenensis (Jacq.) J.F.Macbr. MALPHIGIACEAE Hiptage myrtifolia A.Gray E MALVACEAE *Hibiscus rosa-sinensis L. Hibiscus tiliaceus L. *Malvastrum coromandelianum (L.) Garcke *Sida rhombifolia L. Thespesia populnea (L.) Sol. ex Correa *Urena lobata L. MELASTOMATACEAE Astronidium cf. parviflorum A.Gray E *Clidemia hirta (L.) Don Melastoma denticulatum Labill. Memecylon vitiense A.Gray MELIACEAE Aglaia cf. greenwoodii A.C. Sm. E Dysoxylum sp. E Dysoxylum richii (A.Gray) C.DC. Vavaea amicorum Benth. Vavaea harveyi Seem. E Xylocarpus moluccensis (Lam.) M.Roem. MIMOSACEAE Acacia richii A.Gray E *Adenanthera pavonina L. Entada phaseoloides (L.) Merr. *Leucaena leucocephala (Lam.) de Wit *Mimosa pudica L. Serianthes melanesica Fosberg var. melanesica MORACEAE *Artocarpus altilis (Park.) Fosberg Ficus barclayana (Miq.) Summerh. E Ficus fulvo-pilosa Summerh. E Ficus obliqua Forst. Ficus prolixa G.Forst. Ficus theophrastoides Seem. Ficus vitiensis Seem. Malaisia scandens (Lour.) Planch. Strebulus pendulinus (Endl.) F.v.Muell. MYRISTICACEAE Myristica castaneifolia A.Gray E Myristica gillespieana A.C.Sm. E MYRSINACEAE Maesa corylifolia A.Gray E NYCTAGINACEAE *Bougainvillea ×buttiana Holttum & Standl. *Bougainvillea glabra Choisy OLACEAE Anacolosa lutea Gillespie OLEACEAE Jasminum didymum G.Forst. subsp. didymum Jasminum simplicifolium G.Forst. ONAGRACEAE *Ludwigia octovalvis (Jacq.) Raven PIPERACEAE Macropiper puberulum f. glabrum (DC.) A.C.Sm. PITTOSPORACEAE Pittosporum arborescens A.Gray Pittosporum rhytidocarpum A.Gray E POLYGALACEAE *Polygala paniculata L. RHAMNACEAE Alphitonia franguloides A.Gray E Alphitonia zizyphoides (Spreng.) A.Gray Colubrina asiatica (L.) Brongn. Gouania richii A.Gray E Smythea lanceata (Tul.) Summerh. Ventilago vitiensis A.Gray RHIZOPHORACEAE Bruguiera gymnorrhiza (L.) Lam.f. Crossostylis harveyi Benth. E Crossostylis richii (A.Gray) A.C.Sm. E Rhizophora mangle L. Rhizophora × selala (Salvoza) Toml. Rhizophora stylosa Griff. RUBIACEAE Airosperma vanuense S.Darwin E Antirhea incospicua (Seem.) Christoph. *Gardenia augusta (L.) Merr. Gardenia gordonii Baker E Gardenia hillii Horne ex Baker E Gynochtodes epiphytica (Rech.) A.C.Sm. & S.Darwin *Ixora coccinea L. Ixora elegans Gillespie E *Ixora finlaysoniana Lam. Keppel et al.—Mesic forest of Dogotuki, Vanua Levu Ixora cf. myrtifolia A.C.Sm. E Mastixidendron flavidum (Seem.) A.C.Sm. E Morinda citrifolia L. Morinda myrtifolia A.Gray E Ophiorrhiza leptantha A.Gray Pelagodendron vitiense Seem. E Psychotria amoena A.C.Sm. Psychotria argantha A.C.Sm. E Psychotria gibbsiae S.Moore E Psychotria hypargyraea A.Gray E Psychotria tephrosantha A.Gray E Psychotria sp. (aff. P. macrocalyx A.Gray) E Psydrax odorata (G.Forst.) A.C.Sm. & S.Darwin *Spermacoce assurgens Ruiz & Pavon Tarenna seemanniana A.C.Sm. & S.Darwin E RUTACEAE *Citrus sinensis (L.) Osbeck *Euodia hortensis J.R. & G.Forst. Micromelum minutum (G.Forst.) Seem. Sacromelicope petiolaris (A.Gray) A.C.Sm. E SANTALACEAE Exocarpos vitiensis A.C.Sm. E SAPINDACEAE Dodonaea viscosa (L.) Jacq. Elattostachys falcata (A.Gray) Radlk. SAPOTACEAE Manilkara dissecta (L.f.) Dubard Manilkara smithiana H.J.Lam & Maas Geest. E Palaquium fidjiense Pierre ex Dubard E Palaquium porphyreum A.C.Sm. & S.Darwin E Pouteria cf. garberi (Christophers.) Baehni Pouteria grayana (H.St.John) Fosberg Pouteria membranacea (L.J.Lam) Baehni SIMAROUBACEAE Amaroria soulameoides A.Gray E STERCULIACEAE Commersonia bartramia (L.) Merr. Firmania diversifolia Marsili E Heritiera ornithocephala Kosterm. Melochia grayana A.C.Sm. E THYMELAEACEAE Phaleria glabra (Turrill) Domke Phaleria montana (Seem.) Gilg E Wikstroemia foetida var. vitiensis A.Gray TILIACEAE Grewia crenata (J.R. & G.Forst.) Schinz & Guill. Trichospermum richii (A.Gray) Seem. ULMACEAE Gironniera celtidifolia Gaud. URTICACEAE Leucosyke corymbulosa (Wedd.) Wedd. VERBENACEAE Clerodendrum inerme L. Faradaya ovalifolia (A.Gray) Seem. E Gmelina vitiensis (Seem.) A.. Sm. E 291 *Lantana camara var. aculeata (L.) Moldenke Premna protrusa A.C.Sm. & S.Darwin E *Stachytarpheta urticaefolia (Salisb.) Sims Vitex trifolia var. bicolor Moldenke MONOCOTYLEDONAE (Monocotyledons) AGAVACEAE *Aloe vera L. Cordyline fruticosa (L.) Kunth ARACEAE *Colocasia esculenta L. Epipremnum pinnatum (L.) Engl. *Syngonium podophyllum Schott ARECACEAE Cocos nucifera L. Veitchia filifera (Wendl.) H.E.Moore E BROMELIACEAE *Ananas comosus (L.) Merr. CANNACEAE *Canna indica L. CYPERACEAE Carex dietrichiae Boeck. *Cyperus haspan L. Eleocharis dulcis (Burm.f.) Trin ex Henschel *Eleocharis geniculata (L.) Roem. & Schult. Fimbrystylis dichotoma (L.) Vahl. Gahnia aspera (R.Br.) Spreng. Hypolytrum nemorum subsp. vitiense (C.B.Clarke) T.Koyama Machaerina falcata (Nees) T.Koyama Mariscus javanicus (Houtt.) Merr. & Metcalfe Rhynchospora corymbosa (L.) Britton Scleria lithosperma (L.) Sw. Scleria polycarpa Boeck. DIOSCOREACEAE Dioscorea alata L. Dioscorea nummularia Lam. FLAGELLARIACEAE Flagellaria gigantea Hook.f. Flagellaria indica L. MUSACEAE *Musa (AAA or AAB Group) Simmonds (banana) *Musa (AB Group) (lady’s finger banana) *Musa (AAB) Group (plantain) ORCHIDACEAE Appendicula reflexa Bl. Bulbophyllum gracillimum (Rolfe) Rolfe Bulbophyllum cf. hassallii Kores E Bulbophyllum rotriceps Reichenb.f. Dendrobium catillare Reichenb.f. E Dendrobium platygastrium Reichenb.f. Dendrobium tokai Reichenb.f. ex Seem. Luisia teretifolia Gaud. Malaxis sp. Oberonia heliophila Reichenb.f. Pseuderia smithiana Schweinf. E 292 Spathoglottis pacifica Reichenb.f. Spathoglottis plicata Bl. PANDANACEAE Freycinetia cf. impavida (Hombr. & Jacq.) Stone Pandanus tectorius Warb. POACEAE *Arundo donax L. *Brachiaria mutica (Forssk.) Stapf Centosteca lappacea (L.) Desv. *Dichanthium caricosm (L.) A.Camus Digitaria setigera Roth Garnotia linearis Swallen E Leptaspis angustifolia Summerh. & C.E.Hubb. E Miscanthus floridulus (Labill.) Warb. New Zealand Journal of Botany, 2006, Vol. 44 Oplismenus hirtellus (L.) P.Beauv. *Paspalum conjugatum Berg. *Pennisetum polystachyon (L.) Schult. Saccharum edule L. *Saccharum officinarum L. *Sporobolus cf. diander (Retz.) Beauv. Sporobolus virginicus (L.) Kunth SMILACEAE Smilax vitiensis (Seem.) A.DC. TACCACEAE Tacca leontopetaloides (L.) O.Ktze. ZINGIBERACEAE Alpinia boia Seem. E Alpinia vitiensis Seem.