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Home , Atoll, Island Conservation, Takuu Atoll

Atolls of the Tropical Pacific : Wetlands Under Threat

R. R. Thaman

Contents Introduction ...... 2 Pacific Described ...... 3 Distribution ...... 5 Biodiversity ...... 6 Ecosystem and Habitat Diversity ...... 7 Species and Taxonomic Diversity ...... 10 Genetic Diversity ...... 13 Ecosystem Goods and Services ...... 13 Threats and Future Challenges ...... 13 Conservation of Atoll Ecosystems ...... 19 Conclusion ...... 22 References ...... 23

Abstract Atolls are small, geographically isolated, resource-poor scattered over vast expanses of ocean. There is little potential for modern economic or com- mercial development, and most Pacific atoll countries and communities depend almost entirely on their limited biodiversity inheritances for ecological, economic, and cultural survival in a rapidly globalizing world. Atolls rarely have elevations over 2 or 3 m above level and commonly have extensive areas of intertidal flats, mangroves, shallow , reefs, and limited areas of brackish water marshes or landlocked fossil lagoons and are subject to periodic tidal inundation during extreme weather and tidal events, such as “king ”. Under the Ramsar Convention definition, atolls and their nearshore waters are essentially “wetlands.” Although arguably among the Earth’s “biodiversity

R.R. Thaman (*) The University of the South Pacific, Suva, Fiji e-mail: [email protected]; [email protected]

# Springer Science+Business Media Dordrecht 2016 1 C.M. Finlayson et al. (eds.), The Wetland Book, DOI 10.1007/978-94-007-6173-5_270-1 2 R.R. Thaman

coolspots” with the poorest and highly threatened terrestrial biodiversity inheri- tances on Earth, they are among the last remaining sanctuaries for extensive, but highly threatened populations of breeding seabirds and coral -associated biodiversity. Fortunately, under Ramsar, UNESCO World Heritage, and other relevant conventions and initiatives, the conservation, restoration and sustainable use of atoll biodiversity and associated ethnobiodiversity (the uses, knowledge, beliefs, management systems, taxonomies and language that traditional and the scientific communities have for biodiversity) is now clearly on the conservation agenda.

Keywords Atolls • Biodiversity cool spots • Ethnobiodiversity • Coastal plants • diversity • Food and livelihood security • Global change • Lagoons • Mangroves • Marine biodiversity • Pacific Islands • Sea birds

Introduction

Atolls are small, geographically isolated, resource-poor islands scattered over vast expanses of ocean. There is little potential for modern economic or commercial development, and most Pacific Island atoll countries and communities depend almost entirely on their limited biodiversity inheritances for ecological, economic, and cultural survival. Atolls rarely have elevations over 2 or 3 m above and commonly have extensive areas of intertidal flats, mangroves, shallow lagoons, coral reefs, and limited areas of brackish water marshes or landlocked fossil lagoons and are subject to periodic tidal inundation during extreme weather and tidal events, such as “king tides” (Thaman 2008). Under the Ramsar Convention definition, atolls and their nearshore waters are essentially “wetlands.” Atolls are the opposite of “biodiversity hot spots”–areas such as Amazonia, , , and many Pacific Island areas such as , , , , and the Galapagos – which have very high species and ecosystem diversity and high levels of endemism that are under threat of extinction and degradation (Whittaker 1998). Atolls are among the Earth’s “biodi- versity cool spots” because they have few, if any, endemic plants and animals and among the most impoverished and highly threatened terrestrial and freshwater biodiversity inheritances on Earth, with a high proportion of all economically, culturally, and ecologically important terrestrial plants and animals in danger of extirpation (local extinction) (Thaman 1992a, 2008). Although not as impoverished, atoll marine biodiversity is also under threat and in danger of extirpation, especially on inhabited and urbanized atolls. Despite the poverty, fragility, threatened status, and the obligate dependence of atoll peoples on biodiversity, atoll biodiversity has received only limited attention from the international conservation community, which has focused mainly on the Earth’s “biodiversity hot spots” (Thaman 2008). Fortunately, under Ramsar and a number of other conservation initiatives, , the , and a Atolls of the Tropical : Wetlands Under Threat 3 number of other Pacific Island atoll countries have designated atolls or atoll as “conservation areas” or initiated other initiatives to conserve atoll biodiversity. This chapter discusses the nature of atolls, atoll biodiversity, its value to atoll countries and communities, the threats to and conservation status of atoll biodiversity, and some conservation initiatives, including Ramsar initiatives, that have catalyzed the conservation of atoll biodiversity. Although there are atolls elsewhere, such as the in the , the focus is on all atoll nations and atolls and low-lying limestone of the cultural regions of Melanesia, , and in the tropical Pacific Ocean (Fig. 1).

Pacific Atolls Described

The word atoll comes from the Malayalam word atolu or “reef” or atollon, the native name for the Maldives (Newhouse 1980). Adapting the definitions of Bryan (1953) and Wiens (1962), the term “atoll” refers here to all low-lying oceanic limestone reef islands, with or without lagoons, that have formed on barrier reefs or in the shallow lagoons along the coastlines, or encircle long-submerged ancient volcanoes, which are not associated with nearby high islands or continents (Fig. 2). The term “” refers to the individual smaller islands or “motu” (a Polynesian name for reef islets) that are found on the reefs or in the lagoons of the main atoll islands. In other words, “atolls” include both “true atolls,” the islets of which encircle, border, or are found within a , and individual, separate low-lying limestone reef islands that have no lagoon or may have “secondary” or remnant “fossil” lagoons on the actual limestone island or islets (Thaman 2008). Most atolls have maximum elevations below 3–4 m above sea level, although some have limited areas of coral rubble ramparts deposited over time by high storm waves, limestone pinnacles (e.g., the raised limestone pinnacles on Tikehau Atoll in (Fig. 3), or windblown “” that can reach elevations of over 10 m (e.g., Joe’s Hill which attains 13 m on Atoll in Kiribati). Excluded from this definition of atolls are raised limestone islands or “raised atolls” that have average elevations much higher than 5 m, such as the main islands of the Tongatapu, Ha’apai, and Vava’u groups of and Ouvea, Lifou, and Mare in the Loyalty Islands of New Caledonia; raised phosphatic limestone islands, such as Nauru and Banaba (Ocean Island) and Makatea in the ; and barrier reefs and associated islets surrounding high islands or continents, such as the “almost atolls” of the main Chuuk group and in French Polynesia and islets and reef structures with associated islands such as those on the off northeastern Australia. Also excluded from this definition are “sunken atolls” with- out dry land, such as Middleton and Elizabeth Reefs in the off Australia (Thaman 2008). The above definition of atoll is relatively clear for most of the well-studied “atolls” of Polynesia and Micronesia, although the status of many of the small islands included as atolls in the islands of Melanesia is uncertain due to the lack of detailed published information. For example, Bryan (1953) listed the island of 4 ..Thaman R.R.

Fig. 1 Distribution of the main atoll countries and territories in the Pacific Ocean in the cultural regions of Micronesia, Melanesia, and Polynesia (J. Lowry # Rights remain with the author) Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 5

Fig. 2 Wailagi Lala Atoll, Fiji, partially obscured by cloud (Photo credit: R.R. Thaman # Rights remain with the author)

Ouvea in the Loyalty Islands to the east of New Caledonia as the “world’s largest atoll,” although by his own definition, it is clearly not, but rather a large raised limestone island rising to a maximum elevation of 46 m, much more similar to some of the islands of Tonga, mentioned above (Thaman 2008).

Distribution

Globally, the Earth’s contain more than 400 “atolls” composed of thousands of individual islets! Also included under the definition are numerous small isolated individual low-lying limestone reef islands (Bryan 1953). Most are found in the tropics, especially in the tropical Pacific Ocean, where ocean water temperatures are suitable for coral reefs. The main atoll “groups” in the “cultural areas” of Melanesia, Polynesia, and Micronesia (Fig. 1) include the Marshall, Gilbert, Phoenix, and ; the , , and Northern ; and the Tuamotu Archipelago in French Polynesia, which has 77, the most of any single group. There are also atolls in most of the other countries or island groups in the Pacific Islands, with , New Caledonia, Solomon Islands, Fiji, , and the Federated States of Micronesia, all having atolls. Some of the Northwestern , including the Midway Islands, are also atolls. There is also Clipperton Atoll, a French possession located in the Eastern Pacific about 1080 km southwest of Mexico. 6 R.R. Thaman

Fig. 3 Raised coastal karstified limestone pinnacles, Tikehau Atoll (Photo credit: R.R Thaman # Rights remain with the author)

Pacific countries and territories with no reported true “atolls” include , Tonga, Vanuatu, , , Nauru, and Kosrae (Thaman 2008) (Fig. 1). The largest group of atolls outside the Pacific Islands is the Maldives Archipelago in the Indian Ocean to the southwest of .

Atoll Biodiversity

Atoll biodiversity encompasses (1) ecosystem and habitat diversity, including the staggering diversity of different atolls, atoll islets, and lagoon shapes and sizes, (2) species and taxonomic diversity, (3) genetic diversity, and (4) “ethnobiodiversity” (the knowledge, uses, beliefs, resource-use systems and conservation practices, taxonomies, and language that a given society or community, including the modern scientific community, has for its ecosystems, species, taxa, and genetic diversity). Ethnobiodiversity is here considered an integral component of atoll biodiversity because atoll people and their knowledge, traditions, and spirituality are seen as inseparable from their terrestrial, freshwater, and marine ecosystems, rather than separate external entities. This holistic view is embodied in the Melanesia pidgin concepts of kastom/custom or ples/place; the all-encompassing pan-Polynesian concept of land/fonua, fanua, fenua, henua,or‘enua; or the concepts of te aba in Kiribati and bwirej in the Marshall Islands (Thaman 2004a, 2008). Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 7

Ecosystem and Habitat Diversity

In terms of ecosystem diversity, each “atoll,” whether a true atoll with a central lagoon or lagoons and associated islets or a single low-lying reef island, is somewhat unique, with there being a staggering diversity of atoll island and lagoon types and associated smaller individual islets, lagoons, and ecosystem or habitat types. Depending on the size and type of atoll, ecosystem diversity can include coastal inland, coastal littoral, and mangrove forests; scrublands, grasslands, and herblands; brackish swamps, marshes and ponds (Fig. 4), maricultural areas, and freshwater (groundwater) lenses and wells; swamp taro gardens; agroforests; towns, villages, and houseyard gardens; , rocky limestone , terraces, and limestone reef rock; and reefs, seagrass beds, lagoons, open ocean, sea mounts, and ocean floor. Indigenous atoll vegetation is composed almost exclusively of widespread, ocean-dispersed or, less commonly, wind- or bird-dispersed, salt-tolerant pan-Pacific or pan-tropical coastal plants and mangroves. There are no endemic plant species. Relatively undisturbed indigenous inland atoll forest is now absent on most atolls. Although remnants are found on some inhabited atolls, they are almost exclusively on uninhabited atolls and less accessible uninhabited atoll islets. There remains, however, a significant amount of coastal shoreline forest and scrub vegetation in various stages of disturbance on many atolls, again, more commonly found on isolated uninhabited islets, many of which are globally among the world’s most

Fig. 4 Inland coastal marsh, Temaiku, S. , Kiribati (Photo credit: R.R. Thaman # Rights remain with the author) 8 R.R. Thaman

Fig. 5 Sonneratia alba mangrove forest, Atoll, Kiribati (Photo credit: R.R. Thaman # Rights remain with the author) important seabird reserves. On some of the drier atolls, such as Kiritimati which lies in the tropical dry belt, there are also extensive areas of grassland and low open scrubland. It should be stressed that on atolls there is no surface water in the form of rivers and lakes, with the only real surface water being in the form of limited areas of freshwater marshes and brackish ponds that are found on some atolls, such as the extensive system of landlocked hypersaline ponds on Kiritimati (Christmas) Atoll in the Line Islands of Kiribati. Many of these are polluted, affected by saltwater incursion, decreasing in size or being reclaimed, and are under threat as critical “wetland habitats.” Larger wetter atolls, such as Jaluit in the Marshall Islands (399 cm yearÀ1 rainfall), in Tuvalu (338 cm yearÀ1 rainfall), Butaritari in Kiribati (310 cm yearÀ1 rainfall), and Ontong Java in Solomon Islands, have the most extensive areas of most of the major atoll ecosystems, such as mangroves (Fig. 5), saline swamps, inland atoll forest, and areas of coral reef, and the richest species diversity (Thaman 2008). Although mangroves are found on most of the larger true atolls, almost always along protected lagoon shores or in back- basins in brackish ponds, they are normally not found on lagoon-less reef islands and the atolls of the Central Pacific (Thaman 2008). On many atolls, such as on Funafuti in Tuvalu, extensive areas of mangroves have been reclaimed or destroyed, in the Tuvalu case to build airstrips during World War II (Thaman et al. 2012). Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 9

Fig. 6 Giant swamp taro (Cyrtosperma chamissonis) excavated pit cultivation, Funafuti, Tuvalu (Photo credit: R.R. Thaman # Rights remain with the author)

The most widespread vegetation type on most atolls now consists of coconut- dominated agroforests that are dominated almost exclusively by the coconut palm, often on the best soils, but will include other useful indigenous trees, shrubs and other plants, depending of the level of maintenance of former coconut plantings and regeneration of native plants and introduced weedy species over time. Pandanus (Pandanus spp.), breadfruit (Artocarpus altilis and A. mariannensis), and sometimes bananas (Musa spp.) or other useful trees are also planted, sometimes as small tree groves in more favorable sites, usually near villages or residences or wetter sites (Thaman 1990, 2008; Thaman and Whistler 1996). Excavated taro pits are a unique and specialized wetland agricultural ecosystem found in the central parts of the larger islets of many atolls and in and around villages. These pits have been excavated to the level of the freshwater lens through the limestone bedrock to depths of 1.5–3 m. The artificially enriched soils in these pits, known as taro mucks, are fertile, swampy, and very rich in organic material. The main crop planted in taro pits is giant swamp taro (Cyrtosperma chamissonis (Fig. 6), although common taro (Colocasia esculenta), giant taro (Alocasia macrorrhiza), and bananas (Musa cultivars) often using intensive traditional mulching systems are also occasionally planted. Coconut palms, pandanus, bread- fruit, papaya (Carica papaya), native fig(Ficus tinctoria), and other multipurpose plants are also planted in or near pits. Shrubby species are also found on the margins of the pits; and other multipurpose native trees, including Tournefortia argentea, 10 R.R. Thaman

Fig. 7 Fish harvest from traditional fish weirs, Tikehau Atoll, Tuamotus, French Polynesia (Photo credit: R.R. Thaman # Rights remain with the author)

Guettarda speciosa, and Pipturus argentea, the leaves of which are an important component of the fertilizer or mulch, are also present (Thaman 1990, 2008). There is also limited mariculture practiced on atolls. This includes the culture of black-lip pearl oysters (Pinctada margaritifera) in the Northern Cook Islands and Tuamotus; milkfish (Chanos chanos) mariculture in Kiribati on both Tarawa and Kiritimati Atolls; limited brackish water aquaculture of tilapia (Oreochromis spp.) in Kiribati and on some other atolls, although tilapia is seen as a pest and a hindrance to the mariculture of milkfish in Kiribati; and the mariculture of seaweed (Kappaphycus alvarezii) in Kiritimati and a number of other atolls, although ven- tures have often failed (Gillett 1989; Thaman 2008). There has also been the commercial maricultural production of brine shrimp (Artemia salina) on Kiritimati Atoll, beginning in the 1970s, but abandoned in 1978 (Teeb’aki 1993). Also of note is the use of intricate systems of fish weirs or traps on the intertidal and shallow subtidal areas of atolls as a management strategy for nearshore intertidal finfish resources (Fig. 7).

Species and Taxonomic Diversity

There are four generalizations that apply to the Pacific Islands as a whole for species and taxonomic diversity. These are (1) the “western affinity” of most taxa; (2) a diversity gradient or attenuation of diversity from west to east, as one moves away Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 11 from the Western Indo-Pacific source area of most taxa; (3) a gradual elimination of major groups (higher taxa) of plants and animals from west to east; and (4) a range from very high endemism for high isolated islands to virtually no endemism for the smaller, low-lying limestone islands and atolls and sand cays. The “Theory of Island Biogeography” (MacArthur and Wilson 1967) suggests that species diversity on islands is controlled mainly by island area and distance from the source of colonizing propagules, although as stressed by Fosberg (1974) and Stoddart (1992), species richness on atolls is related more to moisture availability, rainfall extremes, and susceptibility to drought, rather than to island size or to the closeness of source areas of colonizing organisms in the Western Pacific. Atolls in the far west, such as Ontong Java, do however have richer floras and marine biotas. For example, some of the most extensive areas of mangroves and freshwater swamps in Kiribati are found on Butaritari, which is the wettest and most westerly atoll in the main Gilbert group (Thaman 2008). In terms of western affinity of atoll taxa, studies show that almost all terrestrial, freshwater, and marine plant and animal taxa on atolls (e.g., ferns, algae, sea grasses, , echinoderms, marine and terrestrial mollusks, insects, birds, bats, etc.) are mainly of Asian or Western Indo-Pacific origin. The decrease in the total number of species, genera, and families with increasing distance from the Western Indo-Pacific is considerable, and smaller and lower islands have fewer species than larger higher islands. This is due to the differential dispersability of the different organisms, some of which were never able to reach the more distant islands and atolls of the Central Pacific. It is also due to the greater habitat diversity on larger islands, the greater chance of extinction or extirpation (local extinction) among smaller populations on atolls and small islands (often due to prolonged drought or other extreme events, such as “king tides” or tropical cyclones and associated saltwater inundation), and the associated lower probability of initial colonization or recolonization after extirpation or extinction. For example, a large number of marine finfish families absent from the more distant islands on the Pacific Plate in the Central Pacific have shorter larval stages and are unable to disperse over the great distances of open water between oceanic islands. Also estuarine and freshwater habitats are limited to high islands, with such species never being found on atolls (Myers 1991). For whatever reasons, regardless of their distance from the Western Indo-Pacific “hot spot,” as “biodiversity cool spots,” atolls have the most limited species diversity for most taxa, although this is not as dramatic in the marine environment. For example, the number of indigenous fern species drops from 230 in Fiji to 215 in Samoa, 150 in the , and to only 9, 6, 5, and 5 species, respectively, in atolls of the Marshall Islands, Tuvalu, Kiribati, and Tikehau Atoll in the Tuamotus. Even among orchids, which are famous for the high dispersability of their very small light seeds, the same attenuation and almost absolute poverty on atolls are reflected, with the number of orchid species dropping from over 3000 for Papua New Guinea, which has one of the richest orchid floras in the world, to 164 for Fiji, 100 for Samoa, only 3 for Hawai’i, and none for the low-lying atolls of Micronesia and Polynesia, although raised limestone islands, such as Makatea, 12 R.R. Thaman

Fig. 8 Surgeonfishes (Acanthuridae) on , Tikehau Atoll, Tuamotu Archipelago, French Polynesia (Photo credit: R.R. Thaman # Rights remain with the author) which is located only a short distance from the atolls of Tikehau and in the Tuamotus, has two orchids. The number of native angiosperm genera drops from 654 in Solomon Islands, to 476 in Fiji, 302 in Samoa, 263 in Tonga, 201 in the Society Islands, and only 57, 56, 50, and 45 for the Marshall Islands, , Tuvalu, and Tikehau Atoll, respectively, in French Polynesia species (Sekhran and Miller 1996;Kores1991;Whistler1992;Wilder1934;Carlquist 1980;Thaman2008). The disparity is not as great in the marine environment, with benthic marine algae (including green, brown, and red algae) species numbers dropping from 1,185 in North Australia, to 520 for all of Micronesia, 336 for New Caledonia, 302 for Fiji, 219 for the Solomon Islands, 151 for Tahiti, 90 for Samoa, 40 for Nauru, and 268 for the Marshall Islands, one of the larger more westerly atoll groups (N’Yeurt and South 1997). The diversity of corals and reef related finfish diversity is also considerable around atolls relative to terrestrial and freshwater diversity, with atolls considered to be among the best dive sites (Fig. 8). Despite its relative poverty, the conservation of atoll species diversity is critical to the heath and sustainable livelihoods of atoll people. This is reflected in the indig- enous knowledge and taxonomy (ethnobiodiversity) that atolls’ peoples have for their biodiversity. For example, the I-Kiribati (people of Kiribati) have local vernac- ular names for approximately 144 small and 115 large finfishes, 20 sharks, 9 rays, 25 eels, 5 whales or dolphins, 5 sea turtles, 6 sea snakes or snakelike animals, 16 seabirds, and 74 marine mollusks (Thaman 2008). Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 13

Genetic Diversity

The very limited species diversity is, however, enriched by the genetic diversity of cultivars of the limited number of crops that can be cultivated on atolls. In Kiribati, for example, there are reportedly over 200 named cultivars of pandanus (Pandanus spp.), over 30 cultivars of giant swamp taro (Cyrtosperma chamissonis), and at least ten named coconut (Cocos nucifera) cultivars (Small 1972), many of which are now threatened or lost.

Ecosystem Goods and Services

The ecosystem goods and services provided by the highly impoverished terrestrial and freshwater and the richer marine biodiversity of atolls constitute a critical ecological and cultural resource. In the case of plants, this is particularly true, virtually all of which have wide cultural utility, and despite the importance of the sea, plants continue to provide the majority of foods, medicines, handicrafts, con- struction materials, and other material and nonmaterial needs and ecosystem services required by atoll peoples ( 1). In terms of cultural utility, 140 species of widespread indigenous coastal littoral and mangrove vascular plants, almost all of which are found on Pacific atolls, were shown to have 75 different purpose/use categories (Thaman 1992b). Collectively 1024 uses were recorded for these 140 species, ranging from no reported uses (two species) to as many as 125 for the coconut if distinct uses are included (e.g., tools with distinct functions). Another 17 species have 20 or more reported uses, and 29 species have at least 7 uses each. Moreover, the list does not include the more strictly ecological services provided by coastal plants listed in Table 1. For example, for the Marshall Islands, studies showed that there were 168 use categories for 37 of 58 indigenous species found there and another 303 uses for 283 introduced species, the majority of which are ornamentals, food plants, or “weeds” (Thaman 2008). It should be noted that although “plants,” including sea grasses and marine algae, are particularly critical to atoll peoples, coral reefs, currents, areas of , waves, tidal flows and flushing, and other components of marine biodiversity also provide critical ecosystem goods and services that are disproportionately more important to atoll people due to the poverty of terrestrial biodiversity.

Threats and Future Challenges

As stressed above, atolls have among the most impoverished and highly threatened biodiversity inheritances on Earth, with a high proportion of all economically, culturally, and ecologically important terrestrial plants and animals in danger of extirpation (Thaman 2008). Atoll communities are clearly on the frontline in the battle against climate change, sea-level rise, extreme weather and tidal events, loss of biodiversity, environmental 14 R.R. Thaman

Table 1 Ecological and cultural goods and services provided by terrestrial, freshwater, and marine biodiversity on atolls (Adapted from Thaman and Clarke 1993) Ecological Shade/sun protection Soil/substrate Animal/plant habitats improvement Protection from waves/ Sand/sediment provision Pollution control tides Erosion control Spawning/breeding Flood/runoff control grounds Oxygenation Moisture regulation Dispersal facilitation Wind protection Wild animal food Weed/disease control Coastal reinforcement Water purification Protection from salt spray Climate regulation maintenance Cultural/economic Timber/wood Brooms Prop or nurse plants Boatbuilding (canoes) Parcelization/wrapping Food crops Housing/shelter Abrasives/sandpaper Wild/emergency foods Fuel/firemaking Illumination/torches Spices/sauces Woodcarving Insulation Drinks/teas/coffees Tools/weapons Decoration Alcoholic beverages Containers Body ornamentation Masticants/gum Cordage/lashing Fiber/fabric Meat tenderizer Caulking Plaited ware Preservatives Fishing equipment Handicrafts Stimulants/narcotics Floats Clothing/hats Medicines Toys Dyes/paints Aphrodisiacs Musical instruments Paint brushes Fertility control Cages/roosts Tannin/preservative Abortifacients Sails Rubber Ritual exchange Baskets Glues/adhesives Magic/sorcery Religious objects Scents/perfume Totems Deodorants Oils/lubricants Subjects of mythology Toothbrush Poisons Recreation/sport Paper/toilet paper Sunscreens Secret meeting sites Religious objects Insect repellents Commercial/export Switch for children Embalming corpses Products Discipline Cosmetics Tourism attractions degradation, invasive alien species (IAS), and economic uncertainty. Particularly impacted has been , located in Papua New Guinea northeast of Bou- gainville, in which, due to rising sea levels, waves have “savagely eroded the coastline” and tidal incursion has contaminated their limited groundwater supplies and taro gardens. As a result, the Polynesian people of Takuu are considered by some authorities to be the first “climate change refugees,” with many having been resettled Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 15

Fig. 9 Takuu Atoll (October 2004), Papua New Guinea, which has been heavily impacted by sea-level rise (Photo credit: R.R. Thaman # Rights remain with the author) on Bougainville and elsewhere in PNG with predictions being made that “the sea’s relentless advance will extinguish the atoll’s ability to sustain life within the next 2–3 years” (Wane 2005) (Fig. 9). Atoll biodiversity and associated ethnobiodiversity are also threatened by increasing population, urbanization, modern education, commer- cialization, and overexploitation. The highly threatened status of atoll biodiversity is an accelerating phenomenon that began long before European contact when the early Pacific Islanders severely deforested their islands and brought many birds and other species, including some shellfish to extinction or extirpation throughout many areas of Polynesia and Micro- nesia (Kirch 1982; Steadman 1995). Losses are greatest on more urbanized and densely inhabited atolls. Although biodiversity is also threatened on larger high volcanic and raised limestone islands, the need for protection is far greater on atolls where there is far less ecosystem and biotic diversity; smaller, genetically less diverse populations; and higher human population densities (Thaman 2008). Surveys of plants used for specific purposes in Kiribati, Tuvalu, and the Marshall Islands show that there is widespread concern over the loss or scarcity of a wide range of trees and plants used for housebuilding, woodcarving, medicine, body ornamentation, and sacred and other purposes. Even common fruit trees, such as coconut palm and pandanus cultivars, papaya, and breadfruit are reportedly increas- ingly scarce or disappearing; and traditional cultivars of important tree crops and taros are rare or are no longer cultivated because of overemphasis on monoculture, 16 R.R. Thaman

Fig. 10 Frigatebird (Fregata minor) with chick, Ngaon te Taake reserve, Kiritimati Atoll, Kiribati (Photo credit: R.R. Thaman # Rights remain with the author) diseases, tropical cyclones, drought, failure to replant, lack of planting material, and increasing dependence on or taste for imported foods (Thaman and Whistler 1996). Results from studies in the Marshall Islands showed that 38 of a total 61 medicinal plants were considered to be rare or in short supply in some areas by 26 local experts (Taafaki et al. 2006). The loss of medicinal plant biodiversity on atolls is a serious problem because few if any modern medicines are available in rural villages and urban communities. A large number of indigenous birds, reptiles (e.g., lizards and turtles), and a number of terrestrial invertebrates, such as land crabs, are also threatened on atolls. Land crabs, especially the (Birgus latro) and larger land hermit crabs (Coenobita spp.), are now extirpated or rare on most inhabited atoll islets (Thaman 1999a, 2008). Land birds most often mentioned as being rare or endangered include larger land birds such as doves, pigeons, and lorikeets, which are found on some atolls. Also seriously endangered are seabirds and migratory birds that used to be abundant throughout the atoll Pacific. These include noddies, terns, tattlers, god- wits, plovers, frigate birds, boobies, tropic birds, petrels, and shearwaters (Fig. 10). Many of these birds and their eggs, which are important traditional foods and considered delicacies, are now rare or endangered on most atolls. Some species, such as frigate birds and noddies, are of critical importance to atoll peoples as a sign to fishermen of the location and identification of schools of fish (Thaman 2008); and studies in the early 1990s for the 1992 Rio Summit report identified the loss of seabirds as the second most serious environmental concern in Tuvalu (Thaman and Neemia 1991). Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 17

Almost all indigenous birds, the focus of many Ramsar wetland conservation programs, should be given some form of protected status on atolls and their preferred habitats, tree groves, remaining areas of coastal, mangrove, and inland forest and uninhabited islets given protected status. Of particular concern, especially on atolls, has been the widespread removal of groves of Pisonia grandis, the preferred rookery species for a wide range of sea birds. In many cases Pisonia groves have been cleared, beginning in the early colonial period, for the expansion of copra plantations onto the fertile, guano-rich soils resulting from the thousands of generations of seabirds that had occupied the groves prior to their removal (Thaman 2008). Although forest removal, habitat degradation, and hunting have historically been the main causes of the loss of birds, land crabs, and reptiles, the introduction of invasive alien species (IAS), such as pigs, goats, rats, dogs, cats, and ants, may now be the main drivers of the loss of a wide range of birds, crabs and other invertebrates, natural vegetation, farming systems, and human health. The situation is most serious on Guam (which, although not an atoll, has regular air connections with atoll countries) where the accidental introduction of the brown tree snake (Boiga irregularis) near the end of World War II has led to the near extinction of almost all indigenous land birds and the serious endangerment of flying foxes and lizards (Rodda and Fritts 1993; Quammen 1996). In Kiribati, Tuvalu, Tokelau, the Tuamotus, the atolls of Palau and New Caledonia, and possibly other areas, intro- duced ants, such as the yellow crazy ant (Anoplolepis gracilipes) and the little fire ant (Wasmannia auropunctata), both considered to be among the world’s 100 worst IAS, threaten indigenous birds, invertebrates, crops, and human health (CGAPS c. 1996; Wetterer et al. 1997). Although not currently affected by the brown tree snake and some of the more destructive ant species, atoll countries must strengthen their quarantine services to ensure that these pests are not introduced or do not spread to outer islands or islets. There are some serious exotic weeds on atolls that have outcompeted culturally and important indigenous plants. Of particular concern is wedelia or trailing daisy (Sphagneticola/Wedelia trilobata), an introduced ornamental ground cover of trop- ical American origin considered one of the world’s 100 worst . Wedelia has spread out of control on most of the main inhabited atoll islets of Kiribati, Tuvalu, Marshall Islands, and Tikehau Atoll in the Tuamotus, where it is encroaching on interior marshland habitats and coastal herblands. Despite concerted attempts, it has been almost impossible to eradicate due to its ability to spread vegetatively, often via discarded cuttings (Thaman 1999b, 2009). Nutrient pollution from human waste (defecation on the reef is a common practice), sewerage, and pigpen waste, all of which are discharged near the sea or close to the freshwater table, also constitutes a serious threat to the health of atoll wetlands, irrigated taro patches, coral reefs, lagoons, and humans on atolls. Note that there are no larger grazing animals, such as cows, horses, and goats on Pacific atolls because of lack of water, space, and fodder (Thaman 2008). Many nearshore marine species are overfishedforexportorlocalsale. Because of low cash incomes and scarcity of foreign exchange on atolls coupled with the increasing market demand for many marine products, there has been an 18 R.R. Thaman increasing pressure to market shark fin, bêche-de-mer (sea cucumbers), giant clams, large coral reef fish, aquarium fish, live coral, and a number of other marine products, many of which are endangered and listed on the IUCN Red List or on the CITES list of restricted exports (Thaman 2004b). Marine vertebrates that are increasingly threatened include whales and dolphins, marine turtles, sharks and rays, eels, a number of smaller reef and lagoon fish, and a number of larger, commercially important finfish species commonly targeted by spearfishers, hook-and-line fishers, deepwater line fishers, and the live-fish export market, the most common being large rock cods, coral trout, or groupers (Epinephelidae, Serranidae); large parrotfishes (Scaridae); large wrasses (Labridae), particularly the humphead wrasse (Cheilinus undulatus); and treval- lies (Carangidae) (Thaman 2008). Many culturally important mollusks are also rare or extirpated on most inhabited atolls, the most overexploited of which include giant clams (Tridacna spp.), turban snails (Turbo spp.), black-lip pearl oysters (Pinctada margaritifera), conches (Lambis and Strombus spp.), triton trumpet shell (Charonia tritonis), ark shells (Anadara spp.), topshells or trochus (Trochus and Tectus spp.), Venus clams (Periglypta spp.), and mussels (Mytilidae). Crustaceans, including reef and crabs (Calappidae, Carpilidae, Grapsidae, Ocipodidae, Portunidae, and Xanthidae) and mantis shrimps (Stomatopoda), are also rare due to overexploitation. Their declining yields constitute a serious nutritional and economic problem as they are one of the most easily accessible nutritional and commercial resources for low-income coastal communities. For some of these species, there is a need for a total ban on exploitation, whereas for others, there is a need for the establishment of local reserves or commercial or seasonal protection, until such times as stocks recover. Holothurians or bêche-de-mer are overexploited in most atoll countries due to pressure in the past 30 years to export to Asian markets. All species should be given some protected status until stocks recover, with some species being reserved for local consumption or limited local sale. Other considered to be threatened or overharvested in some areas include lobsters, octopus, and squid, all of which are in need of some form of protection, at least the designation of some local marine reserves or the enforcement of seasonal or size restrictions on their exploitation (Thaman 2008). Finally, and most worrying, is that like biodiversity itself, atoll ethnobio- diversity is highly threatened. Many of the current generation, schooled in the modern educational system and living in the cash economy in urbanized, highly populated overexploited areas, often know few of the traditional uses and services provided by atoll biodiversity, let alone the local vernacular names of plants and animals and the places they live. This loss of knowledge has undoubtedly contrib- uted to a loss of appreciation for, and is indirectly associated with, the degradation and loss of biodiversity on atolls (Thaman 2008). The conservation and re-enrichment of this knowledge must be an integral part of attempts at atoll biodiversity conservation. Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 19

Conservation of Atoll Ecosystems

Although the predominant focus of biodiversity conservation has been on endemic, charismatic, and officially in biodiversity hot spots, there is an increasing recognition of the uniqueness, fragility, and need for conservation of atoll biodiversity and other biodiversity cool spots, where seriously threatened very limited biodiversity inheritances constitute the foundation for the sustainability of local communities. This has resulted in an increasing range of collaborative initia- tives under Ramsar, the UNESCO Man and the Biosphere Programme, World Heritage Convention, and other initiatives that have focused on the conservation, restoration, and sustainable use of atoll wetland ecosystems and their surrounding marine areas. Firstly, three Ramsar Wetlands of International Significance have been designated on atolls in the Pacific. These are Jaluit and Namdrik Atolls in the Marshall Islands and Nooto Islet, which is part of North Tarawa Atoll, in Kiribati. Inscripted in 2004, Jaluit is a large atoll covering an area of 69,000 ha comprised of some 91 islets with a land area of 700 ha enclosing a large lagoon. It has relatively pristine areas of coral reefs, intertidal flats, seagrass beds, and mangroves and is an important nesting, nursery, or spawning site for sea turtles, seabirds, and a wide range of finfish and marine invertebrates. About 2000 local residents practice a reasonably sustainable subsistence lifestyle, although some of resources are under threat due to overharvesting, in the case of marine resources for off-island sale (http://www.ramsar.org/jaluit-atoll-conservation-area). Inscripted in 2012, Namdrik Atoll is located about 390 km southwest from the capital atoll of . It has an area of 1,119 ha consisting of two wooded islets with an extensive reef flat lying between them and is unusual because there are no navigable waterways into the central lagoon. Being relatively isolated, the atoll is in near pristine condition, supports an extensive mangrove forest, is home to some 150 species of fish, and supports breeding populations of the endangered hawksbill (Eretmochelys imbricata) and green turtles (Chelonia mydas). The wetland provides many resources for local people, although unsustainable harvesting practices place considerable pressure on the atoll’s unique biodiversity (http://www.ramsar.org/ namdrik-atoll). Inscripted in 2013, Nooto is a relatively pristine northern islet of Tarawa Atoll, the capital atoll of Kiribati. It has an area of 1,033 ha and a wide range of coastal habitats including coral reefs, an extensive lagoon, intertidal flats, mangroves, and nesting, spawning, and nursery sites for sea turtles, bonefish (Albula sp.), and other threat- ened organisms. It is also an important resource island for local North Tarawa Atoll communities (Thaman et al. 1995). Noteworthy international atoll conservation initiatives under the UNESCO Man and the Biosphere (MAB) Programme include the designation in 1977 of Taiaro Atoll in the Tuamotus 540 km to the northeast of Tahiti, as the first atoll ecosystem under MAB, and its redesignation in 2006, after close consultation with local communities, as the Tuamotu Biosphere Reserve to encompass seven additional 20 R.R. Thaman atolls, including Fakarava, the largest and most economically developed atoll; Aratika, Kauehi, Raraka, and Toau Atolls that are inhabited and with navigable passages to interior lagoons; and Taiaro and Niau, two uninhabited closed atolls with interior lagoons but no navigable passages (www.unesco.org/mabdb/br/brdir/direc tory/biores.asp?; Jacques-Bourgeat 2015). In 2007, Ant Atoll, the only uninhabited atoll in of the Federated States of Micronesia, was designated as a MAB Reserve and as an official state protected area in 2010 in efforts to rehabilitate the island, control unsustainable fishing, and protect seabird and coconut crab populations and turtle nesting sites (CSP 2007; Cohen 2015). In 2006 the Papahānaumokuākea Marine National Monument (PMNM) (origi- nally Northwestern Hawaiian Islands Marine National Monument) was established and subsequently inscribed as a World Heritage Site in 2008. PMNM is one of the largest marine protected areas in the world, encompassing an area of 362,000 km2, including ten islands and atolls of the Northwestern Hawaiian Islands. It is interna- tionally recognized for its cultural and natural significance and supports 7,000 marine species, one quarter of which are endemic to the Hawaiian Islands. The islands and shallow water environments are important habitats for threatened species such as the green turtle and the Hawaiian monk seal (Monachus schauinslandi), one of the rarest marine mammals in the world, as well as the 14 million seabirds representing 22 species that breed and nest there. The area has four species of bird found nowhere else in the world, including the world’s most critically endangered duck, the Laysan duck (Anas laysanensis), Laysan albatross (Phoebastria immutabilis), Nihoa millerbird (Acrocephalus familiaris), and Nihoa finches (Telespiza ultima), the latter two which are found only on the small rocky island of Nihoa. Although the Nihoa millerbird used to be present with numbers up to 1,500 in 1915 on Laysan Atoll, it was brought to extinction between 1916 and 1923. There are also interesting species of plants including Pritchardia palms and many arthro- pods and crustaceans. Many marine species seriously overfished in the past are now under protection (PMNM 2014; Birdlife International http://www.birdlife.org/; Morin et al. 1997). In 2007 Kiribati designated the Protected Area (PIPA) as the world’s third largest with an area of 408,250 km2. PIPA includes eight atolls and two submerged coral reefs (PIPA 2007) and has extensive areas of pristine reefs, lagoons, intertidal flats, marshes, seabird, turtle, and land crab nesting areas and rich fisheries resources, many of which are threatened by overexploitation (e.g., illegal fishing) and invasive alien species (IAS), including weeds, rats, cats, rabbits, dogs, and ants, some of which have been the focus of recent successful eradication efforts. These include the successful eradication of Asian rats from McKean Atoll and rabbits from Rawaki Atoll, under the - supported Pacific Invasives Initiative (PII), important for many threatened bird populations which had almost disappeared, and spectacular recovery of the vegeta- tion (Koszler 2010). In 2009, the PIPA was inscribed as a World Heritage Site. In 2008, the New Caledonia barrier reef was formally inscribed as a World Heritage Site under the name The Lagoons of New Caledonia: Reef Diversity and Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 21

Associated Ecosystems. The area includes the central lagoon of Ouvea, a raised limestone island which is sometimes incorrectly referred to as the largest atoll in the world, as well as the Entrecasteaux Atoll group to the north of New Caledonia’s main island (CI 2014). Bikini Atoll Nuclear Test Site in 2010 was inscribed as the first World Heritage Site in the Marshall Islands due mainly to its long history as a nuclear test site and as testimony and memorial to the destruction caused by nuclear testing. The atoll has a total area of 73,500 ha. Alinginae Atoll, another uninhabited atoll, is in the process of applying for inscription as the country’s 2nd World Heritage Site (http://whc.unesco. org/en/list/1339). Other initiatives include the establishment in 2012 by the Cook Islands of the 1.1 million km2 Cook Islands Marine Park, which encompasses the atolls of the North- ern Cook Islands and includes Suwarrow Atoll which has been designated as a Birdlife International Important Bird and Biodiversity Area (IBA) and has just been the focus of a rat eradication program to restore seriously threatened seabird populations (Leahy 2012). And, in 2014 New Caledonia formally created the world’s largest marine park, the Natural Park of the Coral Sea, bringing under management a multiuse marine area of 1.3 million km2, an area rich in atolls, coral reefs, marine mammals, marine turtles, and nesting seabirds (Vorrath 2014). This area includes much of the lagoon of New Caledonia World Heritage Site mentioned above. Almost immediately following the announcement of the estab- lishment of the Natural Park of the Coral Sea, on June 17, 2014, US President Barack Obama used his executive powers to create an even larger marine park in the South- Central Pacific, known as the Pacific Remote Islands Marine National Monument, which protects two million km2 of ocean and the reefs and atolls between Hawaii and from commercial fishing (Neubauer 2014). It must be stressed, however, that the conservation success of all of these large marine protected areas will depend on continued political commitment and the ability to police them. Of particular interest from a wetland conservation perspective have been the continuing conservation initiatives to protect the extensive bird populations on Kiritimati Atoll in Kiribati, the largest true atoll in the world (388 km 2). The bird populations include over 30 species, about 18 of which are seabirds, with the atoll being one of the most important breeding grounds on Earth for some of these species (Streets 1877; Teeb’aki 1993; Thaman et al. 1997) (Fig. 11). These populations have significantly declined and remain seriously threatened by poaching of birds and eggs for food, rats, ants, pigs, and especially feral cats which, since the late nineteenth century, have driven 60 % of the seabirds from the mainland to offshore islets. Initiatives have included the gazetting of the atoll as a bird conservation area by the British in 1960 (when Kiribati was still a colony) and declaring it a wildlife sanctuary in 1975. This included restricting access to the five most important bird nesting areas, the lagoon islets, Cook Island, Motu Tabu, Motu Upua, and Ngaon te Taake, and the sooty tern nesting areas at Southeast Point. The atoll was also the focus of the South Pacific Biodiversity Conservation Programme in the late 1990s, and the conservation of the island is currently carried out by the area by Kiribati national authorities and part of the atoll is being considered as Kiribati’s second Ramsar Site. 22 R.R. Thaman

Fig. 11 Sooty terns (Onychoprion fuscatus) nesting on north , Cook Island (Islet), Kiritimati Atoll, Line Islands, Kiribati (Photo credit: R.R Thaman # Rights remain with the author)

Beginning in 2008 under the New Zealand-funded Pacific Invasives Initiative (PII), IAS were eradicated from 23 of Kiritimati’s lagoon islets, which more than doubled the pest-free area within the lagoon (Koszler 2010). Other initiatives to control IAS on atolls include the eradication of rats from islets on Kayangel Atoll in northern Palau, an area with the biggest population of the Micronesian megapode (Megapodius laperouse), an IUCN red-listed species. This project has been a collaborative effort since 2008 of Birdlife International, PII, and the Palau Conservation Society. The eradication of pigs and rats is the basis for a spectacular recovery of seabird populations on Clipperton Atoll, a French territory to the southwest of Mexico by Island Conservation and partners (Pitman et al. 2005; Birdlife International 2012). On a possibly negative note, Wailagi Lala, Fiji’s only true atoll, which has one of Fiji’s largest colonies of brown noddy (Anous stolidus) and several other seabirds, has been leased to a tourism developer without any provision for the nesting birds (Watling 2013).

Conclusion

In conclusion, the low-lying atolls of the Pacific Ocean and their lagoons and nearshore areas are, under the Ramsar definition, essentially “wetlands.” They are unique wetlands with very limited but highly threatened biodiversity inheritances, most notable of which are extensive populations of breeding seabirds and other birds, for which atolls remain among their most important sanctuaries. As Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 23

“biodiversity cool spots,” the highly threatened terrestrial and nearshore marine biodiversity of atolls constitutes the only biodiversity that atoll nations and peoples have a basis for food and livelihood security and building resilience in the face of global change. Fortunately, under Ramsar, UNESCO World Heritage, and other relevant initiatives, such as the inscription as Ramsar sites of the atolls of Jaluit and Namdrik Atolls in the Marshall Islands and Nooto Islet of North Tarawa Atoll in Kiribati, the conservation and restoration of atoll ecosystems are now firmly on the global conservation agenda.

References

Birdlife International. Eradicating introduced mammals from led to dramatic recovery of seabirds. Presented as part of the BirdLife State of the world’s birds website. 2012. Available from http://www.birdlife.org/datazone/sowb/casestudy/261. Accessed 4 Nov 2014. Bryan Jr EH. Check list of atolls. Atoll Res Bull. 1953;19:1–38. C.I. (Conservation International). New Caledonia: home of the world’s largest marine park. 2014. http://www.conservation.org/projects/Pages/New-Caledonia-Home-of-the-Worlds-Largest- Marine-Park.aspx. Accessed 21 Feb 2015. Carlquist S. Hawaii: a natural history. Kauai/Lawai: Pacific Tropical Botanical Garden; 1980. CGAPS. The silent invasion. Honolulu: Coordinating Group on Alien Pest Species; 1996. Cohen J. Micronesia: keeping Ant Atoll safe. Nature e-News. 2015. http://www.nature.org/ ourinitiatives/regions/asiaandthepacific/micronesia/explore/keeping-ant-safe.xml. Accessed 22 Feb 2015. CSP (Conservation Society of Pohnpei). Ant Atoll declared a UNESCO Biosphere Reserve, vol. 7 (26). Kolonia, Pohnpei: Kaselehlie Press; 2007. p. 12. Fosberg (1974) Proceedings of the Second International Coral Reef Symposium, vol. 1. Brisbane, Australia: The Great Barrier Reef Committee. Gillett R. Tilapia in the Pacific Islands: are there lessons to be learned? Suva: FAO/UNDP Fishery Support Programme; 1989. Jacques-Bourgeat C. Kauehi, un air de paradis. Air Tahiti. 2015;1:14–33. Jaluit Conservation Area: Ramsar sites and countries. http://www.ramsar.org/jaluit-atoll-conserva tion-area. Accessed 4 Nov 2014. Kirch PV. Ecology and adaptation of Polynesian agricultural systems. Archaeol Ocean. 1982;17:1–6. Kores PJ. Family 32. Orchidaceae. In: Smith AC, editor. Flora Vitiensis nova: a new flora of Fiji (spermatophytes only). Comprehensive indices. Lawai/Kauai: Pacific Tropical Botanical Gar- den; 1991. p. 322–576. Koszler N. Restoring biodiversity, one island at a time. Currents. 2010;21:8–13. Leahy S. Tiny Pacific nations create world’s largest marine parks. The Guardian. 2012; 30 August. http://www.theguardian.com/environment/2012/aug/30/pacific-marine-park. Accessed 4 Nov 2014. MacArthur RH, Wilson EO. The theory of island biogeography. Princeton: Princeton University Press; 1967. Morin MS, Conant S, Conant P. Laysan and Nihoa Millerbird (Acrocephalus familiaris). In: Poole A, Gill F, editors. The birds of . Philadelphia and Washington: The Academy of Naural Sciences and The American Ornithologists Union; 1997. No. 302, p. 1–19. Myers RF. Micronesian reef fishes: a practical guide to the coral reef fishes of the tropical central and western Pacific. Barrigada: Coral Graphics; 1991. N’Yeurt ADR, South GR. Biodiversity and biogeography of benthic marine algae in the southwest Pacific, with specific reference to and Fiji. Pac Sci. 1997;51:18–28. 24 R.R. Thaman

Neubauer IL. Is a vast marine sanctuary any use if you can’t police it? Time Magazine. 2014; June 30. http://time.com/2910469/natural-park-of-the-coral-sea-new-caledonia. Accessed 21 Feb 2015. Newhouse J. Marine and terrestrial flora of atolls. South Pac Bull. 1980;30:25–30. Nooto-North Tarawa: Ramsar sites and countries. http://www.ramsar.org/nooto-north-tarawas. Accessed 4 Nov 2014. PIPA (Phoenix Islands Protected Area). 2007. http://www.phoenixislands.org Pitman RL, Balance LT, Bost C. Clipperton Island: pig sty, rat hole and booby prize. Mar Ornithol. 2005;33:193–4. PMNM (Papahānaumokuākea Marine National Monument). 2014. http://www.papahanau mokuakea.gov/wheritage/measures.html. Accessed 4 Feb 2015. Quammen D. The song of the Dodo: Island biogeography in an age of extinction. New York: Touchstone (Simon & Schuster); 1996. Rodda GH, Fritts TH. The brown tree snake on Pacific islands. Pac Sci Assoc Inf Bull. 1993;45 (3–4):1–3. Sekhran N, Miller S. Papua New Guinea country study on biological diversity. Port Moresby: Department of Environment and Conservation; 1996. Small CA. Atoll agriculture in the Gilbert and Ellice Islands. Tarawa: Department of Agriculture; 1972. Steadman DW. Prehistoric extinctions of Pacific Island birds: biodiversity meets zooarchaeology. Science. 1995;267:1123–31. Stoddart DR. Biogeography of the tropical Pacific. Pac Sci. 1992;46:276–93. Streets TH. Some account of the natural history of the Fanning Group of islands. Am Nat. 1877;11:65–72. Taafaki IJ, Fowler MK, Thaman RR, editors. Traditional medicine of the Marshall Islands: the women, the plants, the treatments. Suva: IPS Publications, University of the South Pacific; 2006. Teeb’aki K. Republic of Kiribati. In: Scott DA, editor. A directory of wetlands in Oceania. Slimbridge/Kuala Lumpur: International Waterfowl and Wetlands Research Bureau/Asian Wet- land Bureau; 1993. p. 199–228. Thaman RR. Kiribati agroforestry: trees, people and the atoll environment. Atoll Res Bull. 1990;333:1–29. Thaman RR. Vegetation of Nauru and the Gilbert Islands: case studies of poverty, degradation, disturbance and displacement. Pac Sci. 1992a;46:128–58. Thaman RR. Batiri kei Baravi: the ethnobotany of Pacific Island coastal plants. Atoll Res Bull. 1992b;361:1–62. Thaman RR. Pacific Island biodiversity on the eve of the 21st century: current status and challenges for its conservation and sustainable use. Pac Sci Assoc Inf Bull. 1999a;51(1–4):1–37. Thaman RR. Wedelia trilobata: Daisy invader of the Pacific Islands, IAS Technical Report 99/2. Suva: Institute of Applied Science, University of the South Pacific; 1999b. Thaman RR. Sustaining culture and biodiversity in Pacific Islands with local and indigenous knowledge. Pac Ecol. 2004a;7–8:43–8. Thaman RR. Cool spots under threat: the conservation status of atoll biodiversity and ethno- biodiversity in the Pacific Islands. In: Lee KJ, Tsai H-M, editors. Changing islands – changing worlds: proceedings of Islands of the world VIII. International Small Islands Studies Associa- tion (ISISA); 2004b. p. 60–4. Thaman RR. Atolls – the “biodiversity cool spots” vs “hot spots”: a critical new focus for research and conservation. Micronesica. 2008;40(1/2):33–61. Thaman RR, Fihaki E, Fong T. Plants of Tuvalu: Lākau mo mouku o Tuvalu. Suva: University of the South Pacific, Press; 2012. p. 259. Thaman R. Agrodeforestation and the loss of agrobiodiversity in the Pacific Islands: a call for conservation. In: Jupiter S, Kingsford R, editors. Pacific conservation biology. Special issue on Biodiversity conservation in the Pacific Islands of Oceania, vol. 20(2). 2014. p. 180–92. http:// pcb.murdoch.edu.au Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 25

Thaman RR. Wedelia (Sphagneticola trilobata) – daisy invader of the Pacific Islands: the worst weed in the Pacific? In: Frogier P, Mery P, editors. Proceedings of the 11th Pacific Science Inter- congress and 2nd Symposium on French Research in the Pacific, Papeete/Tahiti; 2009. http:// webistem.com/psi2009/output_directory/cd1/Data/articles/000568.pdf. Accessed 17 Mar 2015. Thaman RR, Clarke WC. Pacific island agroforestry: functional and utilitarian diversity. In: Clarke WC, Thaman RR, editors. Pacific Island agroforestry: systems for sustainability. Tokyo: United Nations University Press; 1993. p. 17–33. Thaman RR, Whistler WA. A review of uses and status of trees and forests in land-use systems in Samoa, Tonga, Kiribati and Tuvalu with recommendations for future action, Working Paper 5, June 1996 (RAS/92/361). Suva: South Pacific Forestry Development Programme; 1996. Thaman RR, Neemia U. Tuvalu: National Report for the United Nations Conference on Environ- ment and Development (UNCED), Rio de Janeiro, June 1992. Office of the Prime Minister, Vaiaku, Funafuti, Tuvalu and South Pacific Regional Environment Programme (SPREP), Noumea; 1991. Thaman RR, Clarke WC, Tebano T, Taniera T, Eritaia B. North Tarawa conservation area: South Pacific biodiversity programme project preparation document. Bikenibeu/Tarawa: Environment Unit, Ministry of Environment and Social Development; 1995. Thaman RR, Tuxson R, Eritaia B. Kiritimati Atoll conservation area: South Pacific biodiversity programme project preparation document. Bikenibeu/Tarawa: Environment Unit, Ministry of Environment and Social Development; 1997. Vorrath S. New Caledonia creates world’s largest marine park. Renew economy; 2014. http:// reneweconomy.com.au/2014/new-caledonia-creates-worlds-largest-marine-park-94028. Accessed 4 Nov 2014. Wane J. Before the flood. Ingenio et Labore: Magazine of the Auckland University; 2005. spring: 10, 12. Watling D. Fiji: state of birds. Suva: NatureFiji-MareqetiViti; 2013. Wetterer JK, Banks PC, Laniawe L, Slotterback JW, Brenner G. Exotic ants at high elevations on Mauna Kea, Hawaii. Pac Sci. 1997;52:228–36. Whistler AW. Vegetation of Samoa and Tonga. Pac Sci. 1992;46:159–78. Whittaker RJ. Island biogeography: ecology, evolution, and conservation. Oxford: Oxford Univer- sity Press; 1998. Wiens HJ. Atoll environment and ecology. New Haven: Yale University Press; 1962. Wilder GP. The flora of makatea, Bulletin 120. Honolulu: Bernice P. Bishop Museum; 1934.