Journal of Marine Sciences

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Ecosystem-Based Management of Pacific Islands

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Volume 2011 |Article ID 358687 |

Jean Kenyon, James Maragos, Douglas Fenner, "The Occurrence of Coral Species Reported as Threatened in Federally Protected Waters of the US Pacific", Journal of Marine Sciences, vol. 2011, Article ID 358687, 12 pages, 2011.

The Occurrence of Coral Species Reported as Threatened in Federally Protected Waters of the US Pacific

Academic Editor: Robert J. Toonen
Received15 Jun 2010
Accepted03 Sep 2010
Published21 Oct 2010


A recent study reported that seventy-five species of reef-building corals, considered to be at elevated extinction risk when assessed by the criteria of the International Union for Conservation of Nature, occur in Pacific waters under United States jurisdiction. Closer examination substantiates records of occurrence for 66 species, while records for the other 9 species were based on misinterpretations or are otherwise uncertain. Of these, at least 55 have been reported from reef habitat under federal protection within National Parks, Marine National Monuments, National Marine Sanctuaries, and National Wildlife Refuges. The highest number of species (31) is found within the Ofu Island unit of the National Park of American Samoa, followed by Kingman Reef (24) and Palmyra Atoll (21), both within the Pacific Remote Islands Marine National Monument. Federally protected areas already in place serve as important habitats for resources whose stewardship needs and priorities may vary over time.

1. Introduction

The International Union for Conservation of Nature (IUCN) Red List of Threatened Species is a tool that is widely used for focusing attention on species of potential conservation concern [1, 2]. Assessments based on the IUCN Red List Criteria allocate species to categories of extinction risk using quantitative rules based on population sizes and decline rates, and range areas and declines. Categories range from “Least Concern” with very little probability of extinction to high risk “Critically Endangered”. Three categories collectively considered “threatened” and representing increasing extinction risk (Vulnerable, Endangered, and Critically Endangered) are intended to serve as one means of setting priority measures for biodiversity conservation [3].

Corals and coral reefs in many regions of the world are reported to be in a state of decline due to numerous local and global anthropogenic stressors including coastal point source pollution, agricultural and land use practices, overutilization for commercial or recreational purposes, and climate change and ocean acidification [49]. In response to their declining condition, three workshops were conducted in 2006 and 2007 to apply the IUCN Red List Categories and Criteria [10] in assessing the extinction risk of all known zooxanthellate reef-building corals from the order Scleractinia plus reef-building octocorals and hydrocorals (families Helioporidae, Tubiporidae, and Milleporidae). Of the 704 species for which there were sufficient data to assign a category, 231 species (32.8%) were assigned to one of the three categories considered threatened [3]. Of these, 75 species were reported to occur in Pacific waters under US jurisdiction (Hawai‘i, American Samoa, Guam, Northern Mariana Islands, Wake Atoll, Johnston Atoll, Howland Island, Baker Island, Jarvis Island, Palmyra Atoll, and Kingman Reef).

Throughout the US Pacific, substantial areas of coral reef habitat are under federal protection within the boundaries of National Wildlife Refuges, Marine National Monuments, National Marine Sanctuaries, or National Parks (Table 1). While the establishment of protected areas may be driven by one or several key resources (e.g., the safeguarding of breeding and calving habitat for the endangered humpback whale by the Hawaiian Humpback Whale National Marine Sanctuary), their legal mandate includes the comprehensive management of the ecological, historical, and aesthetic resources within them. As physical and biological resources continue to respond to environmental change driven by human enterprise, protected areas may provide an umbrella of shelter to habitats and communities not originally envisioned in need of refuge. However, some threats such as climate change are very widespread and challenge the ability of protected areas to limit their effects.

General LocationUnits

Marine National Monuments
 Papahānaumokuākea (PMNM)Northwestern Hawaiian IslandsNihoa , Necker , French Frigate Shoals , Gardner , Maro , Laysan , Lisianski , Pearl and Hermes , Midway NWR, Kure
 Pacific Remote Islands (PRIMNM)Central PacificJohnston Atoll NWR, Howland Island NWR, Baker Island NWR, Jarvis Island NWR, Palmyra Atoll NWR, Kingman Reef NWR, Wake Atoll NWR.
 Rose Atoll (RAMNM)American SamoaRose Atoll NWR
 Marianas Trench, Islands UnitNorthern Mariana IslandsAsuncion, Maug, Farallon de Pajaros
National Marine Sanctuaries
 Hawaiian Islands Humpback Whale   (HIHWNMS) Main Hawaiian IslandsMaui, Lana‘i, Moloka‘i, O‘ahu, Kaua‘i, Hawai‘i
 Fagatele Bay (FBNMS)American SamoaTutuila
National Parks
 NP of American Samoa (NPSA)American SamoaTutuila, Ofu-Olosega, Ta’u
 War in the Pacific NHP (WAPA)Southern Mariana IslandsGuam
 American Memorial Park (AMME)Northern Mariana IslandsSaipan
 Kalaupapa NHP (KALA) Main Hawaiian IslandsMoloka‘i
 Haleakalā NP (HALE) Main Hawaiian IslandsMaui
 Pu‘ukoholā Heiau NHS (PUHE) Main Hawaiian IslandsHawai‘i
 Kaloko-Honokōhau NHP (KAHO) Main Hawaiian IslandsHawai‘i
 Pu‘uhonua o Hōnaunau NHP Main Hawaiian IslandsHawai‘i
  Hawai‘i Volcanoes NP (HAVO) Main Hawaiian IslandsHawai‘i
Other National Wildlife Refuges Ritidian PointGuam

A “unit” is here considered to be a geographically distinct island, atoll, or reef system. NWR: National Wildlife Refuge, : unit within Hawaiian Islands NWR.

The purpose of this paper is to explore the currently known occurrence within federally protected waters of the 75 coral species assessed as threatened by the IUCN and reported by that body to occur within the US Pacific. The number of alleged threatened species occurring in each system of federal protection is reported and those species that are not presently known to occur within the boundaries of US federal protection are identified.

2. Materials and Methods

Assessments based on the IUCN Red List criteria were obtained from Carpenter et al. [3] Supporting Online Material. Peer-reviewed literature, published and unpublished reports, monitoring program databases, and local experts were consulted (Table 2) to corroborate the occurrence of the 75 species stated to occur within US Pacific waters by Carpenter et al. [3] and the IUCN Red List database [43]. Occurrences within the boundaries of federally protected waters were determined from a subset of this suite of sources. No data were available for Haleakalā National Park, Hawai‘i Volcanoes National Park, or American Memorial Park (Brown, pers. comm.).


NPS Pacific Islands Network monitoring database
NOAA Pacific Islands Fisheries Science Center (PIFSC) Coral Reef Ecosystem Division (CRED) database
Pacific Remote Islands[26, 27]
Maragos unpubl. data 2000–2010
Kenyon and Bonito unpubl. data 2005
Rose Atoll[28]
Fenner unpubl. data 2010
Marianas Archipelago[2933]
D. Burdick unpubl. data 2010
G. Paulay, pers. comm.
American Samoa[30, 3442]
Birkeland unpubl. data 1979–2007
Fenner unpubl. data 2010

3. Results

Of the 75 coral species stated to occur in the US Pacific in Carpenter et al. [3] and the IUCN database [43], substantiated records could be identified for 66 species. Six species (Acanthastrea regularis, Acropora tenella, Anacropora puertogalerae, Anacropora spinosa, Pectinia alcicornis, and Physogyra lichtensteini) are shown on distribution maps in Veron [33] as occurring in the Mariana Archipelago, which is the basis of their records in US waters in the IUCN database. However, these records were based on photos submitted to Veron by G. Paulay from Palau and elsewhere that were mistakenly attributed to Guam [31], (G. Paulay pers. comm. Veron pers. comm.), and these 6 species have not been authenticated by other reports from US waters. A seventh species (Acropora lokani) was identified from photographs compared to Veron [33] but was reidentified as A. caroliniana based on the original description of A. lokani in Wallace [44] and the description of A. caroliniana in Wallace [30] (Fenner, pers. comm.). For 2 additional species (Acropora dendrum and Caulastrea echinulata), the initial tentative identifications from photographs are now uncertain (Fenner, pers. comm.).

Of these 66 species, records of occurrence in federally protected waters were identified for 55 species. Forty-three species were recorded within National Parks (NP), 39 species within Marine National Monuments (MNM) and National Wildlife Refuges within the MNM, and 18 within National Marine Sanctuaries (NMS) (Table 3, where data are reported as the number of discrete islands or atolls (“units”) where each species has been recorded. Additionally, a supplementary online table (available at incorporates data presented in Tables , and in a single matrix.) The most frequently occurring species, reported at 10 or more of the 26 major units afforded federal protection from which data have been reported (excluding the Hawaiian Islands Humpback Whale NMS where reported occurrence data were not attributed to the 6 component islands), were Montipora patula (15), Cyphastrea ocellina (13), Montipora flabellata (13), Psammocora stellata (13), Pocillopora elegans (12), Leptoseris incrustans (10), Montipora caliculata (10), and Pocillopora danae (10). Twenty species were recorded from 3 to 8 reef units, and 27 species were recorded at only 1 or 2 reef units.

IUCN Red List CategoryNational ParksMarine National MonumentsNational Marine SanctuariesTotal units

Montipora patulaVU213115
Cyphastrea ocellinaVU211113
Montipora flabellataVU112113
Psammocora stellataVU013113
Pocillopora elegansVU111112
Montipora caliculataVU27110
Pocillopora danaeVU28010
Leptoseris incrustansVU27110
Acropora verweyiVU2608
Acropora acuminataVU1517
Acropora paniculataVU1506
Turbinaria reniformisVU1416
Acropora asperaVU2305
Acropora retusaVU1405
Acropora vaughaniVU1405
Pavona venosaVU3215
Turbinaria stellulataVU1405
Acropora aculeusVU1304
Acropora globicepsVU1304
Acropora microcladosVU1304
Acropora polystomaVU1304
Cyphastrea agassiziVU2214
Heliopora coeruleaVU2204
Acropora palmeraeVU1113
Alveopora verrillianaVU0303
Isopora cuneataVU1203
Montipora dilatataEN0303
Montipora lobulataVU0213
Acropora striataVU1102
Astreopora cucullataVU1102
Isopora crateriformisVU2012
Leptoseris yabeiVU1102
Millepora tuberosaEN2012
Montipora calcareaVU1112
Pavona cactusVU1102
Pavona decussataVU2002
Acanthastrea brevisVU0101
Acropora doneiVU1001
Acropora horridaVU1001
Acropora listeriVU1001
Acropora rudisEN0011
Acropora speciosaVU0101
Alveopora allingiVU1001
Barabattoia laddiVU0101
Euphyllia cristataVU1001
Galaxea astreataVU1001
Montipora angulataVU0101
Montipora australiensisVU1001
Pavona bipartitaVU0101
Pavona diffluensVU1001
Porites horizontalataVU1001
Porites napoporaVU0011
Porites nigrescensVU1001
Porites pukoensisCR0101
Turbinaria mesenterinaVU1001
Acanthastrea hemprichiiVU0000
Acanthastrea ishigakiensisVU0000
Acropora jacquelineaeVU0000
Acropora pharaonisVU0000
Alveopora fenestrataVU0000
Euphyllia paraancoraVU0000
Euphyllia paradivisaVU0000
Millepora foveolataVU0000
Pachyseris rugosaVU0000
Seriatopora aculeataVU0000
Turbinaria peltataVU0000

“Total units” is not strictly additive, as Tutuila hosts both a National Park and a National Marine Sanctuary.

Eleven of the 66 species with substantiated records of occurrence in the US Pacific have not been recorded within federally protected waters (Table 3). Of these, 5 species (Acanthastrea hemprichii, Acanthastrea ishigakiensis, Acropora jacquelineae, Acropora pharaonis, and Euphyllia paradivisa) have been reported from single locations at Tutuila, American Samoa, and were rare, represented by only 1 or 2 colonies (Fenner, pers. comm.). Pachyseris rugosa and Turbinaria peltata have been reported at several locations by several authors in American Samoa [36, 37, 40], (Birkeland unpubl. data, Fenner unpubl. data), indicating that they are relatively more abundant. Four species (Alveopora fenestrata, Euphyllia paraancora, Millepora foveolata, and Seriatopora aculeata) have been reported from Guam [34], (D. Burdick unpubl. data) though abundance data are not available. The occurrence of P. rugosa in the Mariana Archipelago as shown in Veron [33] has not been substantiated by other sources and is based on the misattribution of photographs as indicated previously for 6 other species (Veron pers. comm.).

Of the 39 species recorded within the Marine National Monuments and National Wildlife Refuges (NWR) (Table 4), the highest numbers of Red List threatened coral species have been recorded from Kingman Reef NWR (24) and Palmyra Atoll NWR (21) in the Pacific Remote Islands MNM followed by Rose Atoll NWR and MNM (17). In the Islands Unit of the Mariana Trench MNM, the highest number of Red List threatened coral species has been recorded at Asuncion Island (7). In the Papahānaumokuākea MNM covering the Northwestern Hawaiian Islands, the highest numbers of Red List threatened coral species (5) have been recorded at French Frigate Shoals, Maro Reef, Pearl and Hermes Atoll, and Midway Atoll.

TAXONPacific Remote IslandsRoseMarianasPapahānaumokuākea
JohnstonHowlandBakerJarvisPalmyraKingmanWakeRoseMaugAsuncionFDPNihoaNeckerFFSGardnerMaroLaysanLisinskiP&HMidwayKureNo. units

Montipora patula111111111111113
Psammocora stellata111111111111113
Montipora flabellata11111111111112
Pocillopora elegans1111111111111
Cyphastrea ocellina1111111111111
Pocillopora danae111111118
Leptoseris incrustans11111117
Montipora caliculata11111117
Acropora verweyi111115
Acropora acuminata111115
Acropora paniculata111115
Acropora retusa11114
Acropora vaughani11114
Turbinaria reniformis11114
Turbinaria stellulata11114
Acropora aculeus1113
Acropora aspera1113
Acropora globiceps1113
Acropora microclados1113
Acropora polystoma1113
Alveopora verrilliana1113
Montipora dilatata1113
Cyphastrea agassizi112
Heliopora coerulea112
Isopora cuneata112
Montipora lobulata112
Pavona venosa112
Acanthastrea brevis11
Acropora palmerae11
Acropora speciosa11
Acropora striata11
Astreopora cucullata11
Barabattoia laddi11
Leptoseris yabei11
Montipora angulata11
Montipora calcarea11
Pavona bipartita11
Pavona cactus11
Porites pukoensis11

# Red List species899621245176723354544554

FDP: Farallon de Pajaros; FFS: French Frigate Shoals; P&H: Pearl and Hermes.

Of all the federally protected locations, the highest number of Red List threatened coral species (31) has been recorded from Ofu Island in the National Park of American Samoa (Table 5). Fagatele Bay NMS and the Tutuila section of the National Park of American Samoa each host 12 Red List threatened coral species, despite the substantially smaller size of Fagatele Bay (0.65 km2 and 4.9 km2, resp. )[45, 46]. In Guam, War in the Pacific National Historical Park hosted 6 Red List threatened coral species. In Hawai‘i, the highest number of Red List threatened coral species (5) was recorded in the Hawaiian Islands Humpback Whale NMS.

National ParksMarine Sanctuaries

Leptoseris incrustans111311
Pavona venosa111311
Cyphastrea ocellina111211
Cyphastrea agassizi11211
Isopora crateriformis11211
Millepora tuberosa11211
Montipora patula1111211
Montipora caliculata11211
Turbinaria reniformis1111
Pocillopora elegans1111
Pocillopora danae1120
Pavona decussata1120
Montipora flabellata1111
Montipora calcarea1111
Heliopora coerulea1120
Acropora verweyi1120
Acropora palmerae1111
Acropora aspera1120
Acropora acuminata1111
Turbinaria stellulata110
Turbinaria mesenterina110
Porites nigrescens110
Porites napopora011
Porites horizontalata110
Pavona diffluens110
Pavona cactus110
Montipora lobulata011
Montipora australiensis110
Leptoseris yabei110
Isopora cuneata110
Galaxea astreata110
Euphyllia cristata110
Astreopora cucullata110
Alveopora allingi110
Acropora vaughani110
Acropora striata110
Acropora rudis011
Acropora retusa110
Acropora polystoma110
Acropora paniculata110
Acropora microclados110
Acropora listeri110
Acropora horrida110
Acropora globiceps110
Acropora donei110
Acropora aculeus110

# Red List species632251231125

4. Discussion

This paper summarizes information drawn from publications, reports, museum voucher specimens, monitoring program databases, and coral expert records to the extent that species occurrence could be assigned to benthic habitat within the boundaries of federally protected waters of the US Pacific. However, not all sources provide detailed geographic information; see for example, [32], and not all benthic habitat within the boundaries of federal protection has been surveyed and assessed. Consequently, the extent to which IUCN Red List threatened coral species appear to occur within Marine National Monuments, National Wildlife Refuges, National Parks, and National Marine Sanctuaries of the US Pacific is likely underestimated.

Occurrence records are also likely confounded by problems inherent to current methods of species identification and to classifying corals solely on the basis of morphological criteria. Only relatively recently has coral taxonomy reached a sufficiently comprehensive stage to enable identification of many species over wide areas; see for example, [31, 33, 47]. Corals are highly variable at all scales, and many taxonomic problems remain. A generally accepted hierarchy of data quality, from highest to lowest, is species identification based on examination of voucher specimens, primarily skeletal remains; examination of photographs of both skeletons and living corals; visual field inspection. Even identification of voucher specimens requires resources that are often not available, such as the original description and the type specimen. Nevertheless, many publications and reports are based on visual field identification which for some taxa can be challenging even for veteran experts. In this study, though Pocillopora elegans and P. danae are considered valid species [33], they can be difficult to distinguish from P. meandrina and P. verrucosa, respectively, in the field where they co-occur, and consequently their representation within federally protected waters in the current analysis may be over- or underestimated. Similarly, Porites pukoensis, described from Vaughan [48] and said to be restricted to a single small site off the coast of Moloka‘i Island in Hawai‘i [3, 49], has been reported from Kingman Reef NWR in the Pacific Remote Islands Marine National Monument (Table 4) based on visual identification by a coral taxonomist (Maragos, unpubl. data 2010). Porites pukoensis has also been reported from American Samoa [34, 35], Indonesia [50], Tabuaeran Atoll in the Line Islands and Kanton Atoll in the Phoenix Islands [51, 52], Madagascar [53, 54], Mauritius [54], and Reunion [53, 54], demonstrating the difficulty of validating species distributions even from peer-reviewed publications as the data quality underlying the classification is frequently not stated, and samples that can be used for later verification are rarely collected. Moreover, this species is similar to another species reported from Australia (Porites stephensoni), and molecular analysis may be needed to determine whether these are the same or different species. Montipora dilatata is considered a valid species based on morphological criteria [14, 33, 48, 55] and is thought to have a restricted and disjunct distribution in the Hawaiian Archipelago [3, 13, 14, 33, 56]; however, it has also been reported from Palmyra Atoll NWR [26] and Kingman Reef NWR (Maragos unpubl. data). Moreover, examination of a suite of molecular markers (mitochondrial and nuclear) in addition to a suite of measurements on skeletal microstructure suggest that Montipora dilatata, M. flabellata, and M. cf. turgescens reported from Hawai‘i may be morphological variants of the same biological species [57]. However, Montipora turgescens at its type locality in Australia may not be the same as those identified from Hawai‘i by Veron [33], which are separated by a distance of more than 5000 km, and as a result, the latter may still be a valid species. Occurrence records in this study are based on peer-reviewed literature and the reported observations of scientists widely considered as experts in their field, and therefore, they represent the most current reliable application of coral species identification.

Of the 75 petitioned Pacific species, all but four (Montipora dilatata, M. flabellata, M. patula, and Porites pukoensis) were evaluated under IUCN Red List subcriterion A4, an observed, estimated, inferred, projected, or suspected population size reduction of 50% (for Endangered, EN) or 30% (for Vulnerable, VU) over two generations in the past and one into the future, where the generation length for all but 2 species was considered as 10 years [3]. Species-specific population trend data were not available for the Pacific coral species across their global distributional range, so loss of coral cover within a species distribution in combination with life history traits were used as a surrogate for population reduction. For each species, a quantitative estimate of population reduction was calculated using the coral reef area within its distributional range in conjunction with an estimate of the percent of total coral cover loss or the combined percent of total coral cover loss and critically declining reef [3]. “Total coral cover loss” was operationally defined as the percentage of reef with 90% coral cover loss over at least the past 15 to 20 years, and “critically declining reef” was operationally defined as the percentage of reefs with 50%–90% coral cover loss and likely to join the total coral cover loss category within 10 to 20 years [6]. Three species (Montipora flabellata, M. patula, and Porites pukoensis) were evaluated under IUCN Red List Criterion D, which can be applied to very small populations, where “population” is defined as the total number of individuals of the taxon [10]. Porites pukoensis was listed as Critically Endangered as the population size was estimated to number fewer than 50 mature individuals. Montipora flabellata and M. patula were listed as Vulnerable because they are endemic to Hawai‘i and the number of locations for these species was less than five, where “location” is defined as “a geographically or ecologically distinct area in which a single threatening event can rapidly affect all individuals of the taxon present”. Montipora dilatata was evaluated under IUCN Red List Criterion B, which can be applied to very geographically restricted populations, and was listed as Endangered as its area of occupancy was less than 500 km2 and it existed in less than 5 locations.

While widely viewed as a useful index in estimating extinction risk, the IUCN Red List of Threatened Species carries no weight of law. In October 2009 the nongovernmental organization Center for Biological Diversity petitioned the NOAA to list 83 species of corals under the US Endangered Species Act, including the 75 Pacific species addressed in this study. The Endangered Species Act does have the weight of law. As of this publication, the NMFS is currently leading the process to independently evaluate the extinction risk of these species, and if listed as endangered species, these corals would receive legal protection. Coral reef habitat presently under federal protection harbors at least 55 of the 66 species with validated occurrence records in the US, with three reef units—Palmyra Atoll NWR, Kingman Reef NWR, and the Ofu unit of the National Park of American Samoa—each hosting more than 20 Red List threatened coral species. Protected areas already in place serve as important habitats and may be models for other areas deserving protection because they harbor species at elevated risk.


The authors thank the officers and crew of the NOAA Ship Hi‘ialakai for logistical assistance in conducting dedicated surveys throughout American Samoa and the Pacific Remote Islands Marine National Monument. Eric Brown shared data from the National Park Service Pacific Island Monitoring Network database.

Supplementary Materials

Supplemental Table. Occurrence of 75 IUCN Red Listed Coral Species in the U.S. Pacific and in federally protected areas, and data sources used to substantiate the occurrence records. "1" indicates presence; empty cell indicates no record of occurrence. FDP = Farallon de Pajaros; FFS = French Frigate Shoals; P'H = Pearl and Hermes. National Park and Marine Sanctuary name codes are given in Table 1. 1. VU=vulnerable, EN=endangered, CR=critically endangered. 2. A=supported by existing datasets, B=geographic misattribution, not supported, C=taxonomic misidentification or uncertainly, not supported. 3. Units here represents discrete islands or atolls, except for the HIHWNMS where reported occurrence data were not recorded to individual islands ; units are not strictly additive as Tutuila hosts both a National Park and a Marine Sanctuary, and the island of Hawaii hosts 3 National Parks. 4. Regions covered by unpublished data sources: D. Fenner (American Samoa), J. Maragos (Pacific Remote islands), J. Kenyon (Wake), D. Burdick (Guam), C. Birkeland (American Samoa), National Oceanic and Atmospheric Administration Pacific Islands Fisheries Science Center Coral Reef Ecosystem Division (NOAA PIFSC CRED) database (Marianas), National Park Service Pacific Islands Monitoring Network (NPS PIMN) database (Hawaii National Parks).

  1. Supplementary Table


  1. U. Gärdenfors, C. Hilton-Taylor, G. M. Mace, and J. P. Rodríguez, “The application of IUCN Red List criteria at regional levels,” Conservation Biology, vol. 15, no. 5, pp. 1206–1212, 2001. View at: Publisher Site | Google Scholar
  2. A. S. L. Rodrigues, J. D. Pilgrim, J. F. Lamoreux, M. Hoffmann, and T. M. Brooks, “The value of the IUCN Red List for conservation,” Trends in Ecology and Evolution, vol. 21, no. 2, pp. 71–76, 2006. View at: Publisher Site | Google Scholar
  3. K. E. Carpenter, M. Abrar, G. Aeby et al., “One-third of reef-building corals face elevated extinction risk from climate change and local impacts,” Science, vol. 321, no. 5888, pp. 560–563, 2008. View at: Publisher Site | Google Scholar
  4. G. Hodgson, “A global assessment of human effects on coral reefs,” Marine Pollution Bulletin, vol. 38, no. 5, pp. 345–355, 1999. View at: Publisher Site | Google Scholar
  5. T. A. Gardner, I. M. Côté, J. A. Gill, A. Grant, and A. R. Watkinson, “Long-term region-wide declines in Caribbean corals,” Science, vol. 301, no. 5635, pp. 958–960, 2003. View at: Publisher Site | Google Scholar
  6. C. Wilkinson, Status of Coral Reefs of the World, Australian Institute of Marine Science, Townsville, Australia, 2004.
  7. J. M. Pandolfi, J. B. C. Jackson, N. Baron et al., “Are U.S. coral reefs on the slippery slope to slime?” Science, vol. 307, no. 5716, pp. 1725–1726, 2005. View at: Publisher Site | Google Scholar
  8. J. F. Bruno and E. R. Selig, “Regional decline of coral cover in the Indo-Pacific: timing, extent, and subregional comparisons,” PLoS ONE, vol. 2, no. 8, article e711, 2007. View at: Publisher Site | Google Scholar
  9. J. E. N. Veron, O. Hoegh-Guldberg, T. M. Lenton et al., “The coral reef crisis: the critical importance of < 350 ppm CO2,” Marine Pollution Bulletin, vol. 58, no. 10, pp. 1428–1436, 2009. View at: Publisher Site | Google Scholar
  10. IUCN Red List Categories and Criteria: Version 3.1., 2001,
  11. J. D. Parrish, G. D. Smith, and J. E. Norris, “Resources of the marine waters of Kaloko-Honokohau National Historical Park,” Tech. Rep. 74, Cooperative National Park Resources Study Unit, 1990. View at: Google Scholar
  12. K. Aki, R. Brock, J. Miller et al., “A site characterization study for the Hawaiian Islands Humpback Whale National Marine Sanctuary,” Tech. Rep. 94-06, University of Hawai‘i Sea Grant, 1994. View at: Google Scholar
  13. J. E. Maragos, D. C. Potts, G. Aeby et al., “2000–2002 Rapid Ecological Assessment of Corals (Anthozoa) on shallow reefs of the Northwestern Hawaiian Islands. Part 1: species and distribution,” Pacific Science, vol. 58, no. 2, pp. 211–230, 2004. View at: Google Scholar
  14. D. Fenner, Corals of Hawai‘i, Mutual, Honolulu, Hawai‘i, USA, 2005.
  15. S. Godwin and H. Bollick, “Inventory of intertidal and shallow subtidal marine invertebrates at Kalaupapa National Historical Park,” Tech. Rep. 2006–003, Pacific Cooperative Studies Unit, 2006. View at: Google Scholar
  16. S. E. Kahng and J. E. Maragos, “The deepest, zooxanthellate scleractinian corals in the world?” Coral Reefs, vol. 25, no. 2, p. 254, 2006. View at: Publisher Site | Google Scholar
  17. J. C. Kenyon, P. S. Vroom, K. N. Page, M. J. Dunlap, C. B. Wilkinson, and G. S. Aeby, “Community structure of hermatypic corals at French Frigate Shoals, Northwestern Hawaiian Islands: capacity for resistance and resilience to selective stressors,” Pacific Science, vol. 60, no. 2, pp. 153–175, 2006. View at: Publisher Site | Google Scholar
  18. J. C. Kenyon, M. J. Dunlap, C. B. Wilkinson, K. N. Page, P. S. Vroom, and G. S. Aeby, “Community structure of hermatypic corals at Pearl and Hermes Atoll, Northwestern Hawaiian Islands: unique conservation challenges within the Hawaiian Archipelago,” Atoll Research Bulletin, no. 549, pp. 1–23, 2007. View at: Google Scholar
  19. J. C. Kenyon, C. B. Wilkinson, M. J. Dunlap, G. S. Aeby, and C. Kryss, “Community structure of hermatypic corals at Laysan Island and Lisianski Island/Neva Shoal in the Northwestern Hawaiian Islands: a new layer of scientific exploration,” Atoll Research Bulletin, no. 550, pp. 1–28, 2007. View at: Google Scholar
  20. J. C. Kenyon, M. J. Dunlap, and G. S. Aeby, “Community structure of hermatypic corals at Kure Atoll in the Northwestern Hawaiian Islands: stemming the shifting baseline,” Atoll Research Bulletin, no. 559, pp. 1–25, 2008. View at: Google Scholar
  21. J. C. Kenyon, C. B. Wilkinson, and G. S. Aeby, “Community structure of hermatypic corals at Maro Reef in the Northwestern Hawaiian Islands: a unique open atoll,” Atoll Research Bulletin, no. 558, pp. 1–24, 2008. View at: Google Scholar
  22. J. C. Kenyon, C. B. Wilkinson, and G. S. Aeby, “Community structure of hermatypic corals at Midway Atoll in the Northwestern Hawaiian Islands: a legacy of human disturbance,” Atoll Research Bulletin, no. 581, pp. 1–24, 2010. View at: Google Scholar
  23. D. Minton and M. Carnevale, “A preliminary survey of the marine and intertidal invertebrates of Kalaupapa National Historical Park (Moloka‘i, Hawai‘i),” in Benthic Marine Community Vital Sign Monitoring Protocol—Pacific Island Network, E. Brown, D. Minton, R. Daniel et al., Eds., National Park Service, Oakland, Calif, USA, 2006, Natural Resource Report NPS/PWR/PACN/NRTR-2006/001. View at: Google Scholar
  24. J. Beets, E. Brown, and A. Friedlander, “Inventory of marine vertebrate species and fish-habitat utilization patterns in coastal waters off four national parks in Hawai‘i,” Pacific Cooperative Studies Unit Technical Report 168, Department of Botany, University of Hawai‘i at Mānoa, Honolulu, Hawai‘i, USA, 2010. View at: Google Scholar
  25. P. S. Vroom and C. L. Braun, “Benthic composition of a healthy subtropical reef: baseline species-level cover, with an emphasis on algae, in the Northwestern Hawaiian Islands,” PLoS ONE, vol. 5, no. 3, article e9733, 2010. View at: Google Scholar
  26. G. J. Williams, J. E. Maragos, and S. K. Davy, “Characterization of the coral communities at palmyraatoll in the remote central pacific ocean,” Atoll Research Bulletin, no. 557, pp. 1–32, 2008. View at: Google Scholar
  27. J. E. Maragos and P. L. Jokiel, “Reef corals of Johnston Atoll: one of the world's most isolated reefs,” Coral Reefs, vol. 4, no. 3, pp. 141–150, 1986. View at: Publisher Site | Google Scholar
  28. J. C. Kenyon, J. E. Maragos, and S. Cooper, “Characterization of coral communities at Rose Atoll, American Samoa,” Atoll Research Bulletin. In press. View at: Google Scholar
  29. S. S. Amesbury, D. Ginsburg, T. Rongo, L. Kirkendale, and J. Starmer, “War-in-the-Pacific National Historical Park marine biological survey,” Tech. Rep., University of Guam, 1999. View at: Google Scholar
  30. C. Wallace, Staghorn Corals of the World, A Revision of the Genus Acropora, Australian Commonwealth Scientific and Research Organization, Collingwood, Australia, 1999.
  31. J. E. N. Veron, New species described in Corals of the World, vol. 11 of Marine Science Monograph Series, Australian Institute, 2002.
  32. R. H. Randall, “An annotated checklist of hydrozoan and scleractinian corals collected from Guam and other Mariana Islands,” Micronesica, vol. 35, pp. 121–137, 2003. View at: Google Scholar
  33. J. E. N. Veron, Corals of the World, Australian Institute of Marine Science, Townsville, Australia, 2000.
  34. J. E. Hoffmeister, Some Corals from American Samoa and the Fiji Islands, vol. 343, Carnegie Institution of Washington, 1925.
  35. A. E. Lamberts, “An annotated check list of the corals of American Samoa,” Atoll Research Bulletin, vol. 260–272, no. 264, 1983. View at: Google Scholar
  36. J. E. Maragos, C. L. Hunter, and K. Z. Meier, “Reefs and corals observed during the 1991–92 American Samoa coastal resources inventory,” Tech. Rep., American Samoa Department of Marine and Wildlife Resources, Pago Pago, American Samoa, USA, 1994. View at: Google Scholar
  37. C. Mundy, “A quantitative survey of the corals of American Samoa,” Tech. Rep. number, Department of Marine and Wildlife Resources, Pago Pago, American Samoa, USA, 1996. View at: Google Scholar
  38. D. Fisk and C. Birkeland, “Status of coral communities on the volcanic islands of American Samoa, a re-survey of long-term monitoring sites,” Tech. Rep., Department of Marine and Wildlife Resources, Pago Pago, American Samoa, USA, 2002. View at: Google Scholar
  39. S. Coles, P. R. Reath, P. A. Skelton, V. Bonito, R. C. DeFelice, and L. Basch, “Introduced marine species in Pago Pago Harbor, Fagatele Bay and National Park Coast, American Samoa,” Tech. Rep. 26, Bishop Museum, Honolulu, Hawai‘i, USA, 2003. View at: Google Scholar
  40. E. DiDonato, C. Birkeland, and D. Fenner, “A preliminary list of coral species of the National Park of American Samoa,” Tech. Rep. 155, Pacific Cooperative Studies Unit, 2006. View at: Google Scholar
  41. E. R. Lovell and C. McLardy, “Annotated checklist of the CITES-listed corals of Fiji with reference to Vanuatu, Tonga, Samoa and American Samoa,” Tech. Rep. 415, Joint Nature and Conservation Committee, 2008. View at: Google Scholar
  42. National Park Service, “Corals of National Park of American Samoa,” 2009, View at: Google Scholar
  43. IUCN Red List, 2010,
  44. C. C. Wallace, “New species and a new species group of the coral genus Acropora from Indo-Pacific locations,” Invertebrate Taxonomy, vol. 8, pp. 961–988, 1994. View at: Google Scholar
  45. “Fagatele Bay National Marine Sanctuary,” 2010, View at: Google Scholar
  46. National Park Service, “General Management Plan, National Park of American Samoa,” 1997. View at: Google Scholar
  47. B. W. Hoeksema, “Systematics and ecology of mushroom corals (Scleractinia: Fungiidae),” Zoologica Verehandelingen, vol. 254, p. 471, 1989. View at: Google Scholar
  48. T. W. Vaughan, “Recent madreporaria of the Hawaiian Islands and Laysan,” U.S. National Museum Bulletin, vol. 59, p. 427, 1907. View at: Google Scholar
  49. IUCN Red List of Threatened Species, 2010,
  50. T. Tomascik, A. J. Mah, A. Nontji, and M. K. Moosa, The Ecology of the Indonesian Seas, vol. 7, part 1, Periplus Editions, Hong Kong, 1997.
  51. J. E. Maragos, “Reef corals of Fanning Atoll,” Pacific Science, vol. 28, no. 3, pp. 247–255, 1974. View at: Google Scholar
  52. J. E. Maragos and P. L. Jokiel, “Reef corals of Canton Atoll. I. Zoogeography. An Environmental Survey of Canton Atoll Lagoon 1973,” Final Report NUC TP 395, Naval Undersea Center, Hawai‘i Laboratory, Kailua, Hawai‘i, USA, 1976. View at: Google Scholar
  53. G. Faure, “Annotated check list of corals in the Mascarene Archipelago, Indian Ocean,” Atoll Research Bulletin, vol. 203, pp. 1–26, 1977. View at: Google Scholar
  54. C. R. C. Sheppard, “Coral species of the Indian Ocean and adjacent seas: a synonymized compilation and some regional distributional patterns,” Atoll Research Bulletin, vol. 306-311, no. 307, 1987. View at: Google Scholar
  55. J. E. Maragos, “Order Scleractinia,” in Reef and Shore Fauna of Hawai‘i, D. M. Devaney and L. G. Eldredge, Eds., pp. 158–241, Bishop Museum Press, Honolulu, Hawai‘i, USA, 1977. View at: Google Scholar
  56. IUCN Red List of Threatened Species, 2010,
  57. Z. H. Forsman, G. T. Conception, R. D. Haverkort, R. W. Shaw, J. E. Maragos, and R. J. Toonen, “Is Montipora dilatata an endangered coral species or an ecotype? Genes and skeletal microstructure lump seven Hawaiian species into four groups,” Tech. Rep., NOAA Fisheries Pacific Islands Region Protected Resource Division, 2010. View at: Google Scholar

Copyright © 2011 Jean Kenyon et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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