Atolls of the Tropical Pacific Ocean:Wetlands Under Threat

R. R. Thaman

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

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

R.R. Thaman (*)The University of the South Pacific, Suva, Fijie-mail: Thaman_r@usp.ac.fj; randolph.thaman@usp.ac.fj

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

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

KeywordsAtolls • Biodiversity cool spots • Ethnobiodiversity • Coastal plants • Coral reefdiversity • 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 vastexpanses of ocean. There is little potential for modern economic or commercialdevelopment, and most Pacific Island atoll countries and communities dependalmost entirely on their limited biodiversity inheritances for ecological, economic,and cultural survival. Atolls rarely have elevations over 2 or 3 m above sea level andcommonly have extensive areas of intertidal flats, mangroves, shallow lagoons, coralreefs, and limited areas of brackish water marshes or landlocked fossil lagoons andare subject to periodic tidal inundation during extreme weather and tidal events, suchas “king tides” (Thaman 2008). Under the Ramsar Convention definition, atolls andtheir nearshore waters are essentially “wetlands.”

Atolls are the opposite of “biodiversity hot spots” – areas such as Amazonia,Southeast Asia, Australia, and many Pacific Island areas such as New Guinea, NewCaledonia, Solomon Islands, Hawaii, and the Galapagos – which have very highspecies and ecosystem diversity and high levels of endemism that are under threat ofextinction and degradation (Whittaker 1998). Atolls are among the Earth’s “biodi-versity cool spots” because they have few, if any, endemic plants and animals andamong the most impoverished and highly threatened terrestrial and freshwaterbiodiversity inheritances on Earth, with a high proportion of all economically,culturally, and ecologically important terrestrial plants and animals in danger ofextirpation (local extinction) (Thaman 1992a, 2008). Although not as impoverished,atoll marine biodiversity is also under threat and in danger of extirpation, especiallyon inhabited and urbanized atolls.

Despite the poverty, fragility, threatened status, and the obligate dependence ofatoll peoples on biodiversity, atoll biodiversity has received only limited attentionfrom the international conservation community, which has focused mainly on theEarth’s “biodiversity hot spots” (Thaman 2008). Fortunately, under Ramsar and anumber of other conservation initiatives, Kiribati, the Marshall Islands, and a

2 R.R. Thaman

number of other Pacific Island atoll countries have designated atolls or atoll islets as“conservation areas” or initiated other initiatives to conserve atoll biodiversity. Thischapter discusses the nature of atolls, atoll biodiversity, its value to atoll countriesand communities, the threats to and conservation status of atoll biodiversity, andsome conservation initiatives, including Ramsar initiatives, that have catalyzed theconservation of atoll biodiversity. Although there are atolls elsewhere, such as theMaldives in the Indian Ocean, the focus is on all atoll nations and atolls andlow-lying limestone reef islands of the cultural regions of Melanesia, Polynesia,and Micronesia in the tropical Pacific Ocean (Fig. 1).

Pacific Atolls Described

The word atoll comes from the Malayalam word atolu or “reef” or atollon, the nativename for the Maldives Archipelago (Newhouse 1980). Adapting the definitions ofBryan (1953) and Wiens (1962), the term “atoll” refers here to all low-lying oceaniclimestone reef islands, with or without lagoons, that have formed on barrier reefs orin the shallow lagoons along the coastlines, or encircle long-submerged ancientvolcanoes, which are not associated with nearby high islands or continents (Fig. 2).The term “islet” refers to the individual smaller islands or “motu” (a Polynesianname for reef islets) that are found on the reefs or in the lagoons of the main atollislands. In other words, “atolls” include both “true atolls,” the islets of whichencircle, border, or are found within a lagoon, and individual, separate low-lyinglimestone 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, althoughsome have limited areas of coral rubble ramparts deposited over time by high stormwaves, limestone pinnacles (e.g., the raised limestone pinnacles on Tikehau Atoll inFrench Polynesia (Fig. 3), or windblown sand “dunes” that can reach elevations ofover 10 m (e.g., Joe’s Hill which attains 13 m on Kiritimati 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 theTongatapu, Ha’apai, and Vava’u groups of Tonga and Ouvea, Lifou, and Mare in theLoyalty Islands of New Caledonia; raised phosphatic limestone islands, such asNauru and Banaba (Ocean Island) and Makatea in the Tuamotus; and barrier reefsand associated islets surrounding high islands or continents, such as the “almostatolls” of the main Chuuk group and Bora Bora in French Polynesia and islets andreef structures with associated islands such as those on the Great Barrier Reef offnortheastern Australia. Also excluded from this definition are “sunken atolls” with-out dry land, such as Middleton and Elizabeth Reefs in the Coral Sea 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 smallislands included as atolls in the islands of Melanesia is uncertain due to the lack ofdetailed published information. For example, Bryan (1953) listed the island of

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 3

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4 R.R. Thaman

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

Distribution

Globally, the Earth’s oceans contain more than 400 “atolls” composed of thousandsof individual islets! Also included under the definition are numerous small isolatedindividual low-lying limestone reef islands (Bryan 1953). Most are found in thetropics, especially in the tropical Pacific Ocean, where ocean water temperatures aresuitable for coral reefs.

The main atoll “groups” in the “cultural areas” of Melanesia, Polynesia, andMicronesia (Fig. 1) include the Marshall, Gilbert, Phoenix, and Line Islands; theTuvalu, Tokelau, and Northern Cook Islands; and the Tuamotu Archipelago inFrench Polynesia, which has 77, the most of any single group. There are also atollsin most of the other countries or island groups in the Pacific Islands, with Papua NewGuinea, New Caledonia, Solomon Islands, Fiji, Palau, and the Federated States ofMicronesia, all having atolls. Some of the Northwestern Hawaiian Islands, includingthe Midway Islands, are also atolls. There is also Clipperton Atoll, a Frenchpossession located in the Eastern Pacific about 1080 km southwest of Mexico.

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

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 5

Pacific countries and territories with no reported true “atolls” include Samoa, Tonga,Vanuatu, Wallis and Futuna, Niue, Nauru, and Kosrae (Thaman 2008) (Fig. 1). Thelargest group of atolls outside the Pacific Islands is the Maldives Archipelago in theIndian Ocean to the southwest of India.

Atoll Biodiversity

Atoll biodiversity encompasses (1) ecosystem and habitat diversity, including thestaggering 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 modernscientific community, has for its ecosystems, species, taxa, and genetic diversity).Ethnobiodiversity is here considered an integral component of atoll biodiversitybecause atoll people and their knowledge, traditions, and spirituality are seen asinseparable from their terrestrial, freshwater, and marine ecosystems, rather thanseparate external entities. This holistic view is embodied in the Melanesia pidginconcepts of kastom/custom or ples/place; the all-encompassing pan-Polynesianconcept of land/fonua, fanua, fenua, henua, or ‘enua; or the concepts of te aba inKiribati and bwirej in the Marshall Islands (Thaman 2004a, 2008).

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

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Ecosystem and Habitat Diversity

In terms of ecosystem diversity, each “atoll,” whether a true atoll with a centrallagoon or lagoons and associated islets or a single low-lying reef island, is somewhatunique, with there being a staggering diversity of atoll island and lagoon types andassociated smaller individual islets, lagoons, and ecosystem or habitat types.Depending on the size and type of atoll, ecosystem diversity can include coastalinland, 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; beaches, rocky limestone shores, terraces, and limestonereef 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-tolerantpan-Pacific or pan-tropical coastal plants and mangroves. There are no endemicplant species.

Relatively undisturbed indigenous inland atoll forest is now absent on most atolls.Although remnants are found on some inhabited atolls, they are almost exclusivelyon uninhabited atolls and less accessible uninhabited atoll islets. There remains,however, a significant amount of coastal shoreline forest and scrub vegetation invarious stages of disturbance on many atolls, again, more commonly found onisolated uninhabited islets, many of which are globally among the world’s most

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

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 7

important seabird reserves. On some of the drier atolls, such as Kiritimati which liesin the tropical dry belt, there are also extensive areas of grassland and low openscrubland.

It should be stressed that on atolls there is no surface water in the form of riversand lakes, with the only real surface water being in the form of limited areas offreshwater marshes and brackish ponds that are found on some atolls, such as theextensive system of landlocked hypersaline ponds on Kiritimati (Christmas) Atoll inthe Line Islands of Kiribati. Many of these are polluted, affected by saltwaterincursion, 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 year�1 rainfall), Funafuti in Tuvalu (338 cm year�1 rainfall), Butaritari inKiribati (310 cm year�1 rainfall), and Ontong Java in Solomon Islands, have themost 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 richestspecies diversity (Thaman 2008).

Although mangroves are found on most of the larger true atolls, almost alwaysalong protected lagoon shores or in back-beach basins in brackish ponds, they arenormally 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 ofmangroves have been reclaimed or destroyed, in the Tuvalu case to build airstripsduring World War II (Thaman et al. 2012).

Fig. 5 Sonneratia alba mangrove forest, Butaritari Atoll, Kiribati (Photo credit: R.R. Thaman #Rights remain with the author)

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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 andother plants, depending of the level of maintenance of former coconut plantings andregeneration of native plants and introduced weedy species over time. Pandanus(Pandanus spp.), breadfruit (Artocarpus altilis and A. mariannensis), and sometimesbananas (Musa spp.) or other useful trees are also planted, sometimes as small treegroves 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 ecosystemfound in the central parts of the larger islets of many atolls and in and aroundvillages. These pits have been excavated to the level of the freshwater lens throughthe limestone bedrock to depths of 1.5–3 m. The artificially enriched soils in thesepits, known as taro mucks, are fertile, swampy, and very rich in organic material. Themain crop planted in taro pits is giant swamp taro (Cyrtosperma chamissonis(Fig. 6), although common taro (Colocasia esculenta), giant taro (Alocasiamacrorrhiza), and bananas (Musa cultivars) often using intensive traditionalmulching systems are also occasionally planted. Coconut palms, pandanus, bread-fruit, papaya (Carica papaya), native fig (Ficus tinctoria), and other multipurposeplants are also planted in or near pits. Shrubby species are also found on the marginsof the pits; and other multipurpose native trees, including Tournefortia argentea,

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

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 9

Guettarda speciosa, and Pipturus argentea, the leaves of which are an importantcomponent of the fertilizer or mulch, are also present (Thaman 1990, 2008).

There is also limited mariculture practiced on atolls. This includes the culture ofblack-lip pearl oysters (Pinctada margaritifera) in the Northern Cook Islands andTuamotus; milkfish (Chanos chanos) mariculture in Kiribati on both Tarawa andKiritimati Atolls; limited brackish water aquaculture of tilapia (Oreochromis spp.) inKiribati and on some other atolls, although tilapia is seen as a pest and a hindrance tothe 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 thecommercial maricultural production of brine shrimp (Artemia salina) on KiritimatiAtoll, beginning in the 1970s, but abandoned in 1978 (Teeb’aki 1993). Also of noteis the use of intricate systems of fish weirs or traps on the intertidal and shallowsubtidal areas of atolls as a management strategy for nearshore intertidal finfishresources (Fig. 7).

Species and Taxonomic Diversity

There are four generalizations that apply to the Pacific Islands as a whole for speciesand taxonomic diversity. These are (1) the “western affinity” of most taxa; (2) adiversity gradient or attenuation of diversity from west to east, as one moves away

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

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from the Western Indo-Pacific source area of most taxa; (3) a gradual elimination ofmajor groups (higher taxa) of plants and animals from west to east; and (4) a rangefrom very high endemism for high isolated islands to virtually no endemism for thesmaller, low-lying limestone islands and atolls and sand cays. The “Theory of IslandBiogeography” (MacArthur and Wilson 1967) suggests that species diversity onislands is controlled mainly by island area and distance from the source of colonizingpropagules, although as stressed by Fosberg (1974) and Stoddart (1992), speciesrichness on atolls is related more to moisture availability, rainfall extremes, andsusceptibility to drought, rather than to island size or to the closeness of source areasof colonizing organisms in the Western Pacific. Atolls in the far west, such as OntongJava, do however have richer floras and marine biotas. For example, some of themost extensive areas of mangroves and freshwater swamps in Kiribati are found onButaritari, 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,corals, echinoderms, marine and terrestrial mollusks, insects, birds, bats, etc.) aremainly of Asian or Western Indo-Pacific origin.

The decrease in the total number of species, genera, and families with increasingdistance from the Western Indo-Pacific is considerable, and smaller and lowerislands have fewer species than larger higher islands. This is due to the differentialdispersability of the different organisms, some of which were never able to reach themore distant islands and atolls of the Central Pacific. It is also due to the greaterhabitat diversity on larger islands, the greater chance of extinction or extirpation(local extinction) among smaller populations on atolls and small islands (often due toprolonged drought or other extreme events, such as “king tides” or tropical cyclonesand associated saltwater inundation), and the associated lower probability of initialcolonization or recolonization after extirpation or extinction. For example, a largenumber of marine finfish families absent from the more distant islands on the PacificPlate in the Central Pacific have shorter larval stages and are unable to disperse overthe great distances of open water between oceanic islands. Also estuarine andfreshwater habitats are limited to high islands, with such species never beingfound 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 speciesdiversity 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 to215 in Samoa, 150 in the Society Islands, and to only 9, 6, 5, and 5 species,respectively, in atolls of the Marshall Islands, Tuvalu, Kiribati, and Tikehau Atollin the Tuamotus. Even among orchids, which are famous for the high dispersabilityof their very small light seeds, the same attenuation and almost absolute poverty onatolls are reflected, with the number of orchid species dropping from over 3000 forPapua New Guinea, which has one of the richest orchid floras in the world, to164 for Fiji, 100 for Samoa, only 3 for Hawai’i, and none for the low-lying atolls ofMicronesia and Polynesia, although raised limestone islands, such as Makatea,

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 11

which is located only a short distance from the atolls of Tikehau and Rangiroa inthe Tuamotus, has two orchids. The number of native angiosperm genera dropsfrom 654 in Solomon Islands, to 476 in Fiji, 302 in Samoa, 263 in Tonga, 201 inthe Society Islands, and only 57, 56, 50, and 45 for the Marshall Islands, GilbertIslands, Tuvalu, and Tikehau Atoll, respectively, in French Polynesia species(Sekhran and Miller 1996; Kores 1991; Whistler 1992; Wilder 1934; Carlquist1980; Thaman 2008).

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 inNorth 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 forthe Marshall Islands, one of the larger more westerly atoll groups (N’Yeurt andSouth 1997). The diversity of corals and reef related finfish diversity is alsoconsiderable around atolls relative to terrestrial and freshwater diversity, with atollsconsidered to be among the best dive sites (Fig. 8).

Despite its relative poverty, the conservation of atoll species diversity is critical tothe heath and sustainable livelihoods of atoll people. This is reflected in the indig-enous knowledge and taxonomy (ethnobiodiversity) that atolls’ peoples have fortheir 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).

Fig. 8 Surgeonfishes (Acanthuridae) on fringing reef, Tikehau Atoll, Tuamotu Archipelago,French Polynesia (Photo credit: R.R. Thaman # Rights remain with the author)

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Genetic Diversity

The very limited species diversity is, however, enriched by the genetic diversity ofcultivars 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 (Pandanusspp.), over 30 cultivars of giant swamp taro (Cyrtosperma chamissonis), and at leastten named coconut (Cocos nucifera) cultivars (Small 1972), many of which are nowthreatened or lost.

Ecosystem Goods and Services

The ecosystem goods and services provided by the highly impoverished terrestrialand freshwater and the richer marine biodiversity of atolls constitute a criticalecological 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 thesea, plants continue to provide the majority of foods, medicines, handicrafts, con-struction materials, and other material and nonmaterial needs and ecosystem servicesrequired by atoll peoples (Table 1).

In terms of cultural utility, 140 species of widespread indigenous coastal littoraland mangrove vascular plants, almost all of which are found on Pacific atolls, wereshown to have 75 different purpose/use categories (Thaman 1992b). Collectively1024 uses were recorded for these 140 species, ranging from no reported uses (twospecies) to as many as 125 for the coconut if distinct uses are included (e.g., toolswith distinct functions). Another 17 species have 20 or more reported uses, and29 species have at least 7 uses each. Moreover, the list does not include the morestrictly 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 of58 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 upwelling,waves, tidal flows and flushing, and other components of marine biodiversity alsoprovide critical ecosystem goods and services that are disproportionately moreimportant 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 threatenedbiodiversity inheritances on Earth, with a high proportion of all economically,culturally, and ecologically important terrestrial plants and animals in danger ofextirpation (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

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 13

degradation, invasive alien species (IAS), and economic uncertainty. Particularlyimpacted has been Takuu Atoll, located in Papua New Guinea northeast of Bou-gainville, in which, due to rising sea levels, waves have “savagely eroded thecoastline” and tidal incursion has contaminated their limited groundwater suppliesand taro gardens. As a result, the Polynesian people of Takuu are considered by someauthorities to be the first “climate change refugees,” with many having been resettled

Table 1 Ecological and cultural goods and services provided by terrestrial, freshwater, and marinebiodiversity on atolls (Adapted from Thaman and Clarke 1993)

Ecological

Shade/sun protection Soil/substrateimprovement

Animal/plant habitats

Protection from waves/tides

Sand/sediment provision Pollution control

Erosion control Spawning/breedinggrounds

Flood/runoff control

Oxygenation Moisture regulation Dispersal facilitation

Wind protection Wild animal food Weed/disease control

Coastal reinforcement Water purification Protection from saltspray

Climate regulation Current 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

14 R.R. Thaman

on Bougainville and elsewhere in PNG with predictions being made that “the sea’srelentless advance will extinguish the atoll’s ability to sustain life within the next 2–3years” (Wane 2005) (Fig. 9). Atoll biodiversity and associated ethnobiodiversity arealso threatened by increasing population, urbanization, modern education, commer-cialization, and overexploitation.

The highly threatened status of atoll biodiversity is an accelerating phenomenonthat began long before European contact when the early Pacific Islanders severelydeforested their islands and brought many birds and other species, including someshellfish to extinction or extirpation throughout many areas of Polynesia and Micro-nesia (Kirch 1982; Steadman 1995). Losses are greatest on more urbanized anddensely inhabited atolls. Although biodiversity is also threatened on larger highvolcanic and raised limestone islands, the need for protection is far greater on atollswhere there is far less ecosystem and biotic diversity; smaller, genetically lessdiverse populations; and higher human population densities (Thaman 2008).

Surveys of plants used for specific purposes in Kiribati, Tuvalu, and the MarshallIslands show that there is widespread concern over the loss or scarcity of a widerange of trees and plants used for housebuilding, woodcarving, medicine, bodyornamentation, and sacred and other purposes. Even common fruit trees, such ascoconut palm and pandanus cultivars, papaya, and breadfruit are reportedly increas-ingly scarce or disappearing; and traditional cultivars of important tree crops andtaros are rare or are no longer cultivated because of overemphasis on monoculture,

Fig. 9 Takuu Atoll (October 2004), Papua New Guinea, which has been heavily impacted bysea-level rise (Photo credit: R.R. Thaman # Rights remain with the author)

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 15

diseases, tropical cyclones, drought, failure to replant, lack of planting material, andincreasing 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 medicinalplants 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 seriousproblem because few if any modern medicines are available in rural villages andurban communities.

A large number of indigenous birds, reptiles (e.g., lizards and turtles), and anumber of terrestrial invertebrates, such as land crabs, are also threatened on atolls.Land crabs, especially the coconut crab (Birgus latro) and larger land hermit crabs(Coenobita spp.), are now extirpated or rare on most inhabited atoll islets (Thaman1999a, 2008).

Land birds most often mentioned as being rare or endangered include largerland birds such as doves, pigeons, and lorikeets, which are found on some atolls.Also seriously endangered are seabirds and migratory birds that used to beabundant 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 andconsidered 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 asign to fishermen of the location and identification of schools of fish (Thaman2008); and studies in the early 1990s for the 1992 Rio Summit report identified theloss of seabirds as the second most serious environmental concern in Tuvalu(Thaman and Neemia 1991).

Fig. 10 Frigatebird (Fregata minor) with chick, Ngaon te Taake reserve, Kiritimati Atoll, Kiribati(Photo credit: R.R. Thaman # Rights remain with the author)

16 R.R. Thaman

Almost all indigenous birds, the focus of many Ramsar wetland conservationprograms, should be given some form of protected status on atolls and their preferredhabitats, tree groves, remaining areas of coastal, mangrove, and inland forest anduninhabited islets given protected status. Of particular concern, especially on atolls,has been the widespread removal of groves of Pisonia grandis, the preferred rookeryspecies for a wide range of sea birds. In many cases Pisonia groves have beencleared, beginning in the early colonial period, for the expansion of copra plantationsonto the fertile, guano-rich soils resulting from the thousands of generations ofseabirds that had occupied the groves prior to their removal (Thaman 2008).

Although forest removal, habitat degradation, and hunting have historically beenthe main causes of the loss of birds, land crabs, and reptiles, the introduction ofinvasive alien species (IAS), such as pigs, goats, rats, dogs, cats, and ants, may nowbe 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 seriouson Guam (which, although not an atoll, has regular air connections with atollcountries) where the accidental introduction of the brown tree snake (Boigairregularis) near the end of World War II has led to the near extinction of almostall indigenous land birds and the serious endangerment of flying foxes and lizards(Rodda and Fritts 1993; Quammen 1996). In Kiribati, Tuvalu, Tokelau, theTuamotus, 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 worstIAS, threaten indigenous birds, invertebrates, crops, and human health (CGAPS c.1996; Wetterer et al. 1997). Although not currently affected by the brown tree snakeand some of the more destructive ant species, atoll countries must strengthen theirquarantine services to ensure that these pests are not introduced or do not spread toouter islands or islets.

There are some serious exotic weeds on atolls that have outcompeted culturallyand 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 invasive species.Wedelia has spread out of control on most of the main inhabited atoll islets ofKiribati, Tuvalu, Marshall Islands, and Tikehau Atoll in the Tuamotus, where it isencroaching on interior marshland habitats and coastal herblands. Despite concertedattempts, it has been almost impossible to eradicate due to its ability to spreadvegetatively, often via discarded cuttings (Thaman 1999b, 2009).

Nutrient pollution from human waste (defecation on the reef is a commonpractice), sewerage, and pigpen waste, all of which are discharged near the sea orclose to the freshwater table, also constitutes a serious threat to the health of atollwetlands, irrigated taro patches, coral reefs, lagoons, and humans on atolls. Note thatthere are no larger grazing animals, such as cows, horses, and goats on Pacific atollsbecause of lack of water, space, and fodder (Thaman 2008).

Many nearshore marine species are overfished for export or local sale.Because of low cash incomes and scarcity of foreign exchange on atolls coupledwith the increasing market demand for many marine products, there has been an

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 17

increasing pressure to market shark fin, bêche-de-mer (sea cucumbers), giantclams, large coral reef fish, aquarium fish, live coral, and a number of othermarine products, many of which are endangered and listed on the IUCN Red Listor on the CITES list of restricted exports (Thaman 2004b). Marine vertebratesthat are increasingly threatened include whales and dolphins, marine turtles,sharks and rays, eels, a number of smaller reef and lagoon fish, and a numberof larger, commercially important finfish species commonly targeted byspearfishers, hook-and-line fishers, deepwater line fishers, and the live-fish exportmarket, 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 inhabitedatolls, the most overexploited of which include giant clams (Tridacna spp.), turbansnails (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 shorecrabs (Calappidae, Carpilidae, Grapsidae, Ocipodidae, Portunidae, and Xanthidae)and mantis shrimps (Stomatopoda), are also rare due to overexploitation. Theirdeclining yields constitute a serious nutritional and economic problem as they areone of the most easily accessible nutritional and commercial resources forlow-income coastal communities. For some of these species, there is a need for atotal ban on exploitation, whereas for others, there is a need for the establishment oflocal reserves or commercial or seasonal protection, until such times as stocksrecover.

Holothurians or bêche-de-mer are overexploited in most atoll countries due topressure in the past 30 years to export to Asian markets. All species should be givensome protected status until stocks recover, with some species being reserved for localconsumption or limited local sale. Other marine invertebrates considered to bethreatened or overharvested in some areas include lobsters, octopus, and squid, allof which are in need of some form of protection, at least the designation of somelocal marine reserves or the enforcement of seasonal or size restrictions on theirexploitation (Thaman 2008).

Finally, and most worrying, is that like biodiversity itself, atoll ethnobio-diversity is highly threatened. Many of the current generation, schooled in themodern educational system and living in the cash economy in urbanized, highlypopulated overexploited areas, often know few of the traditional uses and servicesprovided by atoll biodiversity, let alone the local vernacular names of plants andanimals 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 degradationand loss of biodiversity on atolls (Thaman 2008). The conservation andre-enrichment of this knowledge must be an integral part of attempts at atollbiodiversity conservation.

18 R.R. Thaman

Conservation of Atoll Ecosystems

Although the predominant focus of biodiversity conservation has been on endemic,charismatic, and officially threatened species in biodiversity hot spots, there is anincreasing recognition of the uniqueness, fragility, and need for conservation of atollbiodiversity and other biodiversity cool spots, where seriously threatened verylimited biodiversity inheritances constitute the foundation for the sustainability oflocal communities. This has resulted in an increasing range of collaborative initia-tives under Ramsar, the UNESCO Man and the Biosphere Programme, WorldHeritage Convention, and other initiatives that have focused on the conservation,restoration, and sustainable use of atoll wetland ecosystems and their surroundingmarine areas.

Firstly, three Ramsar Wetlands of International Significance have been designatedon atolls in the Pacific. These are Jaluit and Namdrik Atolls in the Marshall Islandsand 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 comprisedof some 91 islets with a land area of 700 ha enclosing a large lagoon. It has relativelypristine areas of coral reefs, intertidal flats, seagrass beds, and mangroves and is animportant nesting, nursery, or spawning site for sea turtles, seabirds, and a widerange of finfish and marine invertebrates. About 2000 local residents practice areasonably sustainable subsistence lifestyle, although some of resources are underthreat 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 thecapital atoll of Majuro. It has an area of 1,119 ha consisting of two wooded isletswith an extensive reef flat lying between them and is unusual because there are nonavigable waterways into the central lagoon. Being relatively isolated, the atoll is innear pristine condition, supports an extensive mangrove forest, is home to some150 species of fish, and supports breeding populations of the endangered hawksbill(Eretmochelys imbricata) and green turtles (Chelonia mydas). The wetland providesmany resources for local people, although unsustainable harvesting practices placeconsiderable 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, thecapital atoll of Kiribati. It has an area of 1,033 ha and a wide range of coastal habitatsincluding 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 Atollcommunities (Thaman et al. 1995).

Noteworthy international atoll conservation initiatives under the UNESCO Manand the Biosphere (MAB) Programme include the designation in 1977 of TaiaroAtoll in the Tuamotus 540 km to the northeast of Tahiti, as the first atoll ecosystemunder MAB, and its redesignation in 2006, after close consultation with localcommunities, as the Tuamotu Biosphere Reserve to encompass seven additional

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 19

atolls, including Fakarava, the largest and most economically developed atoll;Aratika, Kauehi, Raraka, and Toau Atolls that are inhabited and with navigablepassages to interior lagoons; and Taiaro and Niau, two uninhabited closed atolls withinterior lagoons but no navigable passages (www.unesco.org/mabdb/br/brdir/directory/biores.asp?; Jacques-Bourgeat 2015).

In 2007, Ant Atoll, the only uninhabited atoll in Pohnpei State of the FederatedStates of Micronesia, was designated as a MAB Reserve and as an official stateprotected area in 2010 in efforts to rehabilitate the island, control unsustainablefishing, 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 establishedand subsequently inscribed as a World Heritage Site in 2008. PMNM is one of thelargest 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 7,000marine species, one quarter of which are endemic to the Hawaiian Islands. Theislands and shallow water environments are important habitats for threatened speciessuch as the green turtle and the Hawaiian monk seal (Monachus schauinslandi), oneof the rarest marine mammals in the world, as well as the 14 million seabirdsrepresenting 22 species that breed and nest there. The area has four species of birdfound nowhere else in the world, including the world’s most critically endangeredduck, the Laysan duck (Anas laysanensis), Laysan albatross (Phoebastriaimmutabilis), Nihoa millerbird (Acrocephalus familiaris), and Nihoa finches(Telespiza ultima), the latter two which are found only on the small rocky island ofNihoa. Although the Nihoa millerbird used to be present with numbers up to 1,500 in1915 on Laysan Atoll, it was brought to extinction between 1916 and 1923. Thereare also interesting species of plants including Pritchardia palms and many arthro-pods and crustaceans. Many marine species seriously overfished in the past are nowunder protection (PMNM 2014; Birdlife International http://www.birdlife.org/;Morin et al. 1997).

In 2007 Kiribati designated the Phoenix Islands Protected Area (PIPA) as theworld’s third largest marine protected area with an area of 408,250 km2. PIPAincludes eight atolls and two submerged coral reefs (PIPA 2007) and has extensiveareas of pristine reefs, lagoons, intertidal flats, marshes, seabird, turtle, and land crabnesting areas and rich fisheries resources, many of which are threatened byoverexploitation (e.g., illegal fishing) and invasive alien species (IAS), includingweeds, rats, cats, rabbits, dogs, and ants, some of which have been the focus of recentsuccessful eradication efforts. These include the successful eradication of Asian ratsfrom McKean Atoll and rabbits from Rawaki Atoll, under the New Zealand-supported Pacific Invasives Initiative (PII), important for many threatened birdpopulations 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 WorldHeritage Site under the name The Lagoons of New Caledonia: Reef Diversity and

20 R.R. Thaman

Associated Ecosystems. The area includes the central lagoon of Ouvea, a raisedlimestone island which is sometimes incorrectly referred to as the largest atoll in theworld, as well as the Entrecasteaux Atoll group to the north of New Caledonia’smain island (CI 2014).

Bikini Atoll Nuclear Test Site in 2010 was inscribed as the first World HeritageSite in the Marshall Islands due mainly to its long history as a nuclear test site and astestimony and memorial to the destruction caused by nuclear testing. The atoll has atotal area of 73,500 ha. Alinginae Atoll, another uninhabited atoll, is in the process ofapplying 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 1.1million km2 Cook Islands Marine Park, which encompasses the atolls of the North-ern Cook Islands and includes Suwarrow Atoll which has been designated as aBirdlife International Important Bird and Biodiversity Area (IBA) and has just beenthe focus of a rat eradication program to restore seriously threatened seabirdpopulations (Leahy 2012). And, in 2014 New Caledonia formally created theworld’s largest marine park, the Natural Park of the Coral Sea, bringing undermanagement 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 Sitementioned above. Almost immediately following the announcement of the estab-lishment of the Natural Park of the Coral Sea, on June 17, 2014, US President BarackObama 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 andAmerican Samoa from commercial fishing (Neubauer 2014). It must be stressed,however, that the conservation success of all of these large marine protected areaswill depend on continued political commitment and the ability to police them.

Of particular interest from a wetland conservation perspective have been thecontinuing conservation initiatives to protect the extensive bird populations onKiritimati Atoll in Kiribati, the largest true atoll in the world (388 km 2). The birdpopulations include over 30 species, about 18 of which are seabirds, with the atollbeing 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 havesignificantly declined and remain seriously threatened by poaching of birds and eggsfor food, rats, ants, pigs, and especially feral cats which, since the late nineteenthcentury, 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 theBritish in 1960 (when Kiribati was still a colony) and declaring it a wildlife sanctuaryin 1975. This included restricting access to the five most important bird nestingareas, 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 theSouth Pacific Biodiversity Conservation Programme in the late 1990s, and theconservation of the island is currently carried out by the area by Kiribati nationalauthorities and part of the atoll is being considered as Kiribati’s second Ramsar Site.

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 21

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 doubledthe pest-free area within the lagoon (Koszler 2010).

Other initiatives to control IAS on atolls include the eradication of rats from isletson Kayangel Atoll in northern Palau, an area with the biggest population of theMicronesian megapode (Megapodius laperouse), an IUCN red-listed species. Thisproject has been a collaborative effort since 2008 of Birdlife International, PII, andthe Palau Conservation Society. The eradication of pigs and rats is the basis for aspectacular recovery of seabird populations on Clipperton Atoll, a French territory tothe 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 ofFiji’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 andnearshore areas are, under the Ramsar definition, essentially “wetlands.” They areunique wetlands with very limited but highly threatened biodiversity inheritances,most notable of which are extensive populations of breeding seabirds and otherbirds, for which atolls remain among their most important sanctuaries. As

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

22 R.R. Thaman

“biodiversity cool spots,” the highly threatened terrestrial and nearshore marinebiodiversity of atolls constitutes the only biodiversity that atoll nations and peopleshave a basis for food and livelihood security and building resilience in the face ofglobal change. Fortunately, under Ramsar, UNESCO World Heritage, and otherrelevant initiatives, such as the inscription as Ramsar sites of the atolls of Jaluitand Namdrik Atolls in the Marshall Islands and Nooto Islet of North Tarawa Atoll inKiribati, the conservation and restoration of atoll ecosystems are now firmly on theglobal conservation agenda.

References

Birdlife International. Eradicating introduced mammals from Clipperton Island led to dramaticrecovery 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. Accessed22 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 FisherySupport 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 Nov2014.

MacArthur RH, Wilson EO. The theory of island biogeography. Princeton: Princeton UniversityPress; 1967.

Morin MS, Conant S, Conant P. Laysan and Nihoa Millerbird (Acrocephalus familiaris). In: PooleA, Gill F, editors. The birds of North America. Philadelphia and Washington: The Academy ofNaural 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 centraland western Pacific. Barrigada: Coral Graphics; 1991.

N’Yeurt ADR, South GR. Biodiversity and biogeography of benthic marine algae in the southwestPacific, with specific reference to Rotuma and Fiji. Pac Sci. 1997;51:18–28.

Atolls of the Tropical Pacific Ocean: Wetlands Under Threat 23

Neubauer IL. Is a vast marine sanctuary any use if you can’t police it? Time Magazine. 2014; June30. http://time.com/2910469/natural-park-of-the-coral-sea-new-caledonia. Accessed 21 Feb2015.

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.orgPitman 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 challengesfor 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 indigenousknowledge. 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 – changingworlds: 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 researchand conservation. Micronesica. 2008;40(1/2):33–61.

Thaman RR, Fihaki E, Fong T. Plants of Tuvalu: Lākau mo mouku o Tuvalu. Suva: University ofthe South Pacific, Press; 2012. p. 259.

Thaman R. Agrodeforestation and the loss of agrobiodiversity in the Pacific Islands: a call forconservation. In: Jupiter S, Kingsford R, editors. Pacific conservation biology. Special issue onBiodiversity conservation in the Pacific Islands of Oceania, vol. 20(2). 2014. p. 180–92. http://pcb.murdoch.edu.au

24 R.R. Thaman

Thaman RR. Wedelia (Sphagneticola trilobata) – daisy invader of the Pacific Islands: the worstweed 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: ClarkeWC, Thaman RR, editors. Pacific Island agroforestry: systems for sustainability. Tokyo: UnitedNations University Press; 1993. p. 17–33.

Thaman RR, Whistler WA. A review of uses and status of trees and forests in land-use systems inSamoa, Tonga, Kiribati and Tuvalu with recommendations for future action, Working Paper5, 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: SouthPacific biodiversity programme project preparation document. Bikenibeu/Tarawa: EnvironmentUnit, Ministry of Environment and Social Development; 1995.

Thaman RR, Tuxson R, Eritaia B. Kiritimati Atoll conservation area: South Pacific biodiversityprogramme project preparation document. Bikenibeu/Tarawa: Environment Unit, Ministry ofEnvironment 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.

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