The State of Coral Reef Ecosystems of the U.S. Pacific Remote Island Areas

Rusty Brainard1, Jim Maragos2, Robert Schroeder1,3, Jean Kenyon1,3, Peter Vroom1,3, Scott Godwin4, Ronald Hoeke1,3,

Areas Greta Aeby5, Russell Moffitt1,3, Marc Lammers1,6, Jamison Gove1,3, Molly Timmers1,3, Stephani Holzwarth1,3, Steve Kolinski7

INTRODUCTION AND SETTING The U.S. Pacific Remote Island Areas (PRIAs) are nine sovereign Federal territories that straddle the equator in the central Pacific. All are single reef ecosystems that are a part of a large central Pacific biogeographic and geological province consisting mostly of ancient low reef islands and atolls (Figure 12.1; Stoddart, 1992). Six of the PRIAs are atolls or atoll reefs: Johnston Atoll (16˚N, 169˚W), Palmyra Atoll (5˚53΄N, 162˚05΄W), Kingman Reef (6˚25΄N, 162˚23΄W), which constitute the three northernmost of the U.S. Line Islands; Rose Atoll (14˚S, 168˚W), the easternmost of the Samoan Islands; Wake Atoll (20˚N, 155˚W), the northernmost of the Marshall Islands; and Midway Atoll (28˚N, 177˚W), near the northwestern end of the Hawaiian Archipelago. The remaining three PRIAs are low reef islands within one degree latitude of the equator: Jarvis Island (00˚S, 160˚W), in the central U.S. Line Islands, Howland Island (00˚18΄S, 160˚01’W); and Baker Island (00˚13΄N, 176˚38΄W), the two northernmost of the U.S. Phoenix Islands. All except Wake and Johnston are National Wildlife Refuges (NWRs) administered by the U.S. Fish and Wildlife Service (USFWS) and all fall under co- jurisdiction of the U.S. Department of Interior (DOI) and the U.S. Department of Commerce (DOC), except Johnston, which is managed by the U.S. Department of Defense, and Palmyra, which is under the joint jurisdiction of DOI, DOC, and The Nature Conservancy. Pacific Remote Island Although all nine are outside the political jurisdiction of other U.S. Pacific States and Territories (Hawaii, American Samoa, Guam, and the Commonwealth of the Northern Mariana Islands [CNMI]), Rose and Midway Atolls are geographically a part of American Samoa and Hawaii, respectively. For organizational purposes, Midway Atoll is mostly treated in the chapter on the Northwestern Hawaiian Islands (NWHI), and Rose Atoll is partially covered in the chapter on American Samoa. Swains Island (14˚S, 168˚W), the northernmost island in American Samoa, is also a remote reef island but it is not technically a PRIA because it falls under the jurisdiction of the Territory of American Samoa.

All of the PRIAs were uninhabited at the time of their discovery by Americans and Europeans over the past two centuries, although Polynesians (and Micronesians, in the case of Wake) probably visited all of the islands periodically over many centuries to harvest fish and wildlife. The U.S. claimed most of the islands via the Guano Act of 1856. Except for Palmyra, Kingman and Rose, the PRIAs lie within arid zones of the tropical Pacific, with insufficient groundwater and rainfall to support continuous human habitation. Moreover, Kingman, lacks vegetated islets, and the land area at Rose is too small and vulnerable to storms to allow habitation. Although Palmyra is certainly capable of supporting human settlements, it is unclear as to why it remained uninhabited during recent centuries. The lack of human habitation allowed the coral reef ecosystems of the PRIAs to remain completely pristine until the early 20th century. Even today all lie beyond the influence of urban centers, associated pollutants, and major shipping lanes.

Most of the PRIAs were materially modified during the World War II (WWII) era: the U.S. constructed and occupied military bases at Johnston, Palmyra, Wake, Midway, and Baker, while Kingman, Jarvis, and Howland were also briefly occupied or utilized during the war era. With the closure of the military base at Johnston in early 2004, only Wake Atoll remains an active U.S. military base. The seven NWRs in the PRIAs were established between 1924-2001, and all are presently no-take island and marine protected areas (MPAs) except Palmyra, on which limited catch and release sport fishing for bonefish and offshore pelagic catch for local consumption are allowed. 1 NOAA Fisheries, Pacific Islands Fisheries Science Center 2 U.S. Fish and Wildlife Service 3 University of Hawaii, Joint Institute for Marine and Atmospheric Research 4 Bishop Museum 5 Department of Land and Natural Resources, Division of Aquatic Resources 6 University of Hawaii, Hawaii Institute of Marine Biology 7 NOAA Fisheries, Pacific Islands Regional Office page 338 Figure 12.1. Locator mapfor thePacificRemoteInsular Areas (PRIAs).Map: A. Shapiro. The StateofCoralReefEcosystemsthePacificRemoteIsland Areas page 339 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Ocean currents transport and distribute larvae among and between different atolls and islands, and particularly in the Pacific equatorial region, define sea surface temperatures (SSTs) and available nutrient regimes. The North Equatorial Current (NEC), Equatorial Counter Current (ECC), Equatorial Undercurrent or Cromwell Current (EUC), and South Equatorial Current (SEC) provide the mechanism by which many species are

Areas distributed among the PRIAs, nearby central Pacific islands, the main Hawaiian Islands (MHI), as well as other distant regions.

ENVIRONMENTAL AND ANTHROPOGENIC STRESSORS

Climate Change and Coral Bleaching Federally sponsored coral reef surveys conducted between 1979-2004 reveal that all PRIAs have experienced coral bleaching during the past 25 years. Pre-2000 observations by Maragos (1979, 1994) include reports and photographs of localized coral bleaching of the ocean reefs at Wake in mid-1979 and mass coral bleaching in progress at Rose off all reefs to depths of 25 m in April 1994. NOAA and USFWS scientists who visited Wake in 1998 did not report any bleaching. Several USFWS visits and surveys of Rose between 1995-1999 and recent surveys of Rose in 2002 and 2004 sponsored by the National Oceanic and Atmospheric Administration (NOAA) reveal that corals are still recovering from a mass bleaching event of 1994. Coral bleaching was also reported at Johnston and Kingman in the 1990s (P. Jokiel, pers. comm.). The four NOAA and USFWS- sponsored expeditions to Baker, Howland, Jarvis, Kingman, and Palmyra between 2000-2004 strongly suggest that mass bleaching occurred within the few years before 2000, and that corals have remained in recovery phase since then. Additional evidence on recent bleaching at Palmyra reveal that the reefs off the broad western reef terrace at Palmyra supported thriving Acropora staghorn coral thickets in 1987 (Maragos, Pacific Remote Island 1988) that died and degenerated to rubble deposits by November 1998 (Molina and Maragos, 1998). The distribution of large Porites heads and numerous small Pocillopora coral heads present on the terrace in late 1998 suggest mass coral bleaching in the mid-1990s rather than coral mortality caused by storm waves at Palmyra.

During 12 benthic surveys at Johnston Atoll in January 2004, mild bleaching of relatively few coral colonies were found at eight of the 12 sites. Mild bleaching was observed on Montipora patula, M. capitata, and Acropora cytherea.

During much of June through December 2002, both remotely sensed and in situ measurements of SST around Jarvis Island, Baker Island, and Howland Island were extremely high, at times 2˚C or more above maximum mean conditions, for prolonged periods (Figure 12.2). Prolonged temperature anomalies such as these have been implicated in most widespread coral bleaching events. Due to their remote nature, these reefs were not surveyed during or immediately after this period of SST increase. Biological monitoring Figure 12.2. Satellite and in situ temperatures at Baker and Jarvis Islands during by experienced scientists in early 2002 and 2003 showing anomalously high SST. Both satellite Pathfinder SST (Baker 2004 did not yield visual evidence – thick blue line, Jarvis – thick green line) and in situ temperatures (z~=-15 m) at that widespread coral mortality had Baker (thin blue line) and Jarvis (thin green line) show values significantly exceeding long-term mean climatological values (Baker–light gray line, Jarvis–dark gray line). occurred in the wake of a possible Coral Reef Watch bleaching threshold of maximum monthly mean SST plus 1°C are bleaching event in 2002. included for reference. Source: Brainard et al., 2004. page 340 any While Keli in1984(Figure12.4). passed the 20 tropical Because 60 knots(Figure12.4). Kingman/Palmyra (EEZs) intensity the and the Mexico which These of In T activities. damselfi and frequently patches having in 2004. around around disease A Howland andBakerIslands assessment ofcoraldisease. at al. ring-like tissue yet Montipora and estimated no. were disease. In Johnston Diseases ropical Storms general, Johnston tropical January good (2001) qualitative degrees been equator PRIAs. of U.S. storms Montipora damage seemed evident the colonies) death are islands The any and of Howland Baker near storm . sh lesion was condition Line found of Surveys Johnston Atoll found impacts No prevalence=0.5%; Howland, most patchy storm the , observed 2004, that Cyclogenesis head latitude. signs corals denuded the ( Atoll in Plectroglyphididon at Johnston to tropical and conducted to and , activity PRIAs Island the assessment (Figure in 92% white U.S. active was similar head reef be events, Island west tissue during the were of surveys Phoenix EEZ. of center atolls with were is Baker estimated of associated Line these As disease. habitats , NWHI of during cyclones located syndrome away on skeleton 12.3). undergo in catching limited since the disease loss. Johnston, few and a However a are found late producing although at and (storm result, , acroporids qualitative sites were tropical Islands or from was six of located colonies Aeby six January 1979 W These summer at associated to Phoenix Jarvis at of Small ork cyclolysis to signs storm were coral surveyed. sites sites sites on a Montipora conducted formation) with Johnston the sp.) , , 3.1%, any Kingman, be higher only NWHI occurrences include et storm to the a diseases in Island PRIAs within the magnitude and systems Islands between 04 Photo:G. Aeby. 2004. 12.3. Figure one table which chapter). events (storm latitude south with experiences early Hurricane The ring The StateofCoralReefEcosystemsthePacificRemoteIsland in to the toward of at Palmyra, coral, have the the these are these is 12 lagoon syndrome, out average and the at death) fall on higher have than west/northwest past located sites also Johnston Montipora patula Montipora T the Acropora of major coral (Figure the ypes three, storms John Jarvis, the due the 60 western been or been a at than NWHI prevalence reef higher of spin years. eastern out eastern to other where Johnston Ekeka in 12.4). diseases Howland, would harsh observed are observed what cytherea 1994, of ecosystems of to Pacific. the f frequency areas, much af of northwards Three from Johnston Atoll with abnormal growths, January growths, abnormal with Johnston Atoll from the in and Pacific has weather Most fected have the path of Hurricane 1992, Atoll included north. , disease western and in been within Montipora more it PRIAs cyclones Additionally storms of the been is corals center to in reached Baker of almost conditions. subjected NWHI. reported Signifi quantify the before the common tropical growth produces Keoni (no. Pacific caused that have PRIAs that Exclusive experience have tissue all cant of wind , One reaching develop due in tropical storm anomalies diseased and travel for abnormal to tropical than Signs been primarily are 1993, hurricanes tropical speeds loss disease a the to characterize greater Economic not events their between the NWHI observed low of cyclones the of syndrome, and storm f colonies/total coral documented, growths other depressions on in the frequencies proximity longitude by that Hurricane degree excess than that Acropora (average extreme centers, coast disease has 10 areas. Zones of in Areas have coral both with and and any the not to of of of of page 341 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

wave energy events. In summary, tropical storms represent a moderately frequent disturbance event experienced by coral reefs

Areas at Johnston Island, potentially influencing physical and biological reef structure, although certainly not to the degree experienced by reefs and terrestrial environments in the western Pacific. Kingman, Palmyra, Jarvis, Howland, and Baker, on the other hand, are rarely, if ever, directly subjected to significant tropical storms and hurricanes.

Significant Wave Events While the direct impacts of tropical storms on the coral reef ecosystems of most of the PRIAs islands and atolls are relatively rare, the impacts of large wave events resulting from distant tropical and extratropical

Pacific Remote Island storms may be significant. Episodic events with wave heights around 2-3 m and periods of 15-20 seconds occur at frequent intervals throughout the year in the equatorial PRIAs (Baker, Howland, Jarvis, Palmyra, and Kingman). Although this is an order of Figure 12.4. A map showing the paths and intensities of tropical storms passing magnitude less than NWHI wintertime near the PRIAs, 1979-2004. Year of storm, storm name and storm strength on the Saffir-Simpson scale (H1-5) are indicated for each. Map: A. Shapiro. Source: NOAA wave events, the equatorial PRIAs are Coastal Services Center. much less seasonal in terms of wave energy and these relatively more moderate events occur throughout the year (Figure 12.5). These events generally approach this area from the northeast through the northwest during the boreal winter, from the southwest in the boreal summer, and 4 to a smaller extent, from the west during the northern hemisphere’s typhoon season. The wave climate of Johnston Atoll is likely very similar to that of the NWHI and MHI, with a

large number of extremely energetic ave Power (W/m) x 10 events in the wintertime and very W consistent, moderate waves in the summer.

These episodic wave events Month subject the shallow-water coral reef communities to a great deal more Figure 12.5. Climatological values of wave power (W/m) derived from NOAA Buoy #51028 located along the equator at 154°W near Christmas Island from 1997-2004. energy than the average energy level. Source: NOAA National Data Buoy Center. As such, wave climate likely plays a page 342 activities promotes Nature activity permanently activities The T be assessedatthistime. and The are nowbeingproposed. eco-toxicological corroding sites of corals including a Coral transfer oftheatolltoUSFWSmanagement. DoD waste, completed contaminants testing, remain, Palmyra Environmental chemicals, for The Coastal Pollution runof have for of central Surface prior totheclosureofmilitarybasethereinearly2004. few by Palmyra There Coastal DevelopmentandRunoff fi fundamental shes, andinvertebratesofthecoralreefecosystemsPRIAs. ourism andRecreation nearby atolls foundations periods collecting cyanobacteria of military pollution status f. to deteriorated generating monitoring 2004 has (Figure the dioxins, Pacific, conduct at especially Conservancy and storage runof is and W periodic at metallic Johnston not the being the .W and exceeding of bases compatible low Palmyra termination at role some rainfall closed f at .II to been 12.6) military is Restoration task atoll. and one and leaving W military reef additional data of dissolved nearly in proposed dieback studies the or ake at debris. at Johnston other burning (Kingman chemical forming several characteristic of Atoll any and islands. lie as Johnston all Palmyra, collected Atoll, Presently atoll a Atoll sponsors destroying contamination behind within debris a non-existent decade sport-fishing coastal proliferation chemical buildings at of ecotourism of source cleanup Additional Program and environment. could iron participating of Acropora and including the The munitions, military Reef) an fuel dump, and Midway abandoned of , coordinated maintaining sites from or development Atoll f The and paved arid of not the all contaminants in Palmyra. of and more, of freshwater lacks funded drums chemical in northwest zone, military , ern coastofBaker Island. ing Figure and facilities the runways research and thus permanent site fuels, Although The StateofCoralReefEcosystemsthePacificRemoteIsland PRIAs. Johnston and at the by biogeographic Johnston BAK-5P 12.6. stockpiling contaminants in Howland . generating agents The coast draining were all have the The at solvents, institutions Evidence W are Nature Most PRIAs near the ake, PRIA land, Atolls of demolished not and was incapable Baker and and the of Johnston been of Midway sewage and and associated of (spatial runways also Conservancy since housed abandoned Agent the and Baker periodic demolition and Source: Maragos andV in numerous removed. all the PRIAs 2000-2004 toxics the of , universities. by have pollution and Orange in and up site Atoll collecting Acropora at W the WWII the fuels to Johnston Palmyra, .W lie at vertical) of porous of several U.S. and late Chemical contaminants. The .II Johnston post-war within military underground defoliants, and showed dieback and era. 1980s, the USFWS rainfall Department During carbonate distributions various are ocean Jarvis, explosives the eit, 2004. thousand USFWS An at high as Agent evidence designed arid atoll but USFWS and dump the is all part Howland, fuel toxic atmospheric The DoD’ many presently zone generating research soils past of that of military Disposal site of in permanent storage of which Defense of and as an of 2003, will of year contaminants corals, characteristic impacts f the catchments the acceptable hazardous and s Defense be urging personnel , station released northwest all tanks System nuclear nuclear surface tropical funded monitor algae, (DoD) Baker Areas but from the at at a - - page 343 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

a) catch-and-release sport-fishing for lagoon bonefish, b) blue water sport-fishing of offshore pelagic species, with catch limited to what can be consumed on the island, c) sport diving, d) snorkeling, e) hiking, f) beach- going, g) touring historic sites, and h) birding. Aside from Palmyra, all other central Pacific NWRs are closed to commercial fishing. Access to any NWR requires a Special Use Permit issued by the NWR manager. Permits

Areas are granted only for actions compatible with the conservation and protection of fish, wildlife, and vegetation inhabiting the refuges. The military facility at Wake Atoll allows sport diving, beach-going, touring historical sites, and other recreational activities for resident DoD workers and some visitors, although Wake is not a tourism destination.

Fishing The islands and surrounding waters of Howland, Baker, and Jarvis; Kingman Reef; and most of Palmyra Atoll are currently NWRs under the management of the USFWS. Fishing and unauthorized entry in these waters are prohibited; however, current levels of poaching are largely unknown. Anecdotal evidence of possible fishing activity includes the presence of fishing lines entangled on the reef >50 m deep, as seen on NOAA’s Pacific Island Fisheries Science Center-Coral Reef Ecosystem Division’s (PIFSC-CRED) towed-optical assessment device video surveys in 2004. The Nature Conservancy owns one-third of Palmyra, primarily for ecotourism (e.g., catch-and-release fishing for bonefish [Albula spp.] in the lagoon). In recent years, occasional trips have been made by Hawaii-based fishing vessels to Kingman and Palmyra for coastal sharks (fining) and bottomfish. There is also recent anecdotal evidence of occasional fishing at Kingman and Palmyra (e.g., Fijian fishers on tuna boats based out of San Francisco). However, it is presently not possible to quantify such effort or catch, nor whether by foreign vessels or passing yachts. Johnston Atoll is under the control of the U.S. military with overlay status as a NWR. Recreational and subsistence fishing is regularly practiced Pacific Remote Island by the temporary workers on the island. Based on a survey during the late 1980s, catch was dominated by soldierfish (Holocentridae). Other fish taken included bigeyes (Priacanthidae), flagtails (Kuliidae), mullet (Mugilidae), goatfish (Mullidae), jacks (Carangidae), parrotfish (Scaridae), and surgeonfish (Acanthuridae). It was also common for island residents to regularly ship coolers of fish and corals to Hawaii; however, this practice has recently been prohibited.

Trade in Coral and Live Reef Species Trade in live reef organisms in the Pacific is primarily for live food fish and ornamental aquarium species, including colorful fish, corals, other invertebrates, and “live rock.” Total value of the trade exceeds $1 billion (USD) per year (Barber and Pratt, 1997; Sadovy and Vincent, 2002). Large species of fish, such as humphead wrasse (Cheilinus undulatus), large groupers (Serranidae) and emperors (Lethrinidae) are captured and ordered live in upscale restaurants, primarily in Southeast Asia (e.g., Hong Kong), where prices can exceed $100 (USD) per kg. The practice can be harmful as sodium cyanide is often used to facilitate capture, resulting in mass mortality of coral reef communities. Targeted large fish are characteristically long-lived and can be rapidly depleted locally. This results in collection operations moving eastward from Southeast Asia in search of more productive reefs (e.g., Micronesia and the Marshall Islands). There is no evidence to date that live reef fish harvesters have reached the PRIAs. Any such take would be illegal as these areas are no-take NWRs, with one (Wake) under control of the U.S. military. However, the targeted species/sizes do occur at these islands, which are remote and largely unprotected due to the lack of enforcement. The long transport distance of live product to Asian markets may render these U.S. islands economically unfeasible for operations, with the possible exception of Wake Atoll, an active, occupied military base.

There is currently no known live reef ornamental fishing operating in the PRIAs. However, some export of live corals was documented in the 1980s from Johnston Atoll to Hawaii, though DoD and USFWS cooperatively eliminated this activity thereafter. The DoD closed its operations at Johnston in 2004, and the atoll is again an uninhabited NWR. As such it is now vulnerable to unauthorized collection by fishers from nearby Hawaii (1,500 km to the northwest) because of the present lack of on-site enforcement and surveillance.

page 344 the The Aquatic InvasiveSpecies cables. seen during storage Having seen duringeachsurveyyear 2002 forereef mining during Long-line material wasdumpedof during present debris 2001, Fishing Marine Debris cyanobacteria thatcandisplacenativespeciesandsmotherreefbuildingorganisms. and from re-introducing of PRIAs and All ships donotcallatanyofthePRIAsexceptperiodicallyPalmyra scheduled to: W All Ships, Boats,andGroundings W cryptogenic 12.1, are have alien are Compiled conducted at1 the PRIAs. Bishop ake. ake Island,an activemilitarybase,are notavailable. the a) the PRIAs PIFSC- U.S. at associated resident aids ef ruptured species and experienced season. NOAA including USFWS 2002, either forts all accumulation except the past. towed-diver PRIAs served Museum, Periodic nets of and Documentation zone Line to three gear f data recorded. are supply CRED W the by navigation. term and oceanographic Johnston disposal .W seabirds. and In Islands, reaching of W vessels rodents are 1 sitesatJohnston vulnerable was northwestern as and Although from survey species the 2004, .II with ake, transports Kingman 2004 refers anthropogenic a both vessels in assault uninhabited the encountered specialists PIFSC- Pacific surveys these 2002 these Documented at Johnston, in one and or years greatest run in and the based Dissolved f thenorthwesternanchorage. name, Howland, to of the W the Palmyra the to and large to Reef. ef on 12 ake marine atolls species the military CRED altered other call research 1970s forts in anchorage density consisted Palmyra; ship U.S. . at T in 2004. over risk arawa native suf except the physical at fishing and Johnston at via Hawaii, objects This reveal alien groundings Atoll bytheBishopMuseum. Atoll. iron fered Line W alien and Baker base items habitats. all of and of U.S. the the ake Palmyra of These vessels mass range sites. Atoll. unknown net, and from for net of species and nets the past 1980s, species of a hulls that Both for , alteration Line to include were have encrusted Jarvis, Atoll. major several Baker was c) with The StateofCoralReefEcosystemsthePacificRemoteIsland offload USFWS refueling Phoenix corroding kills surveys The three encountered (if After marine A unauthorized and are of every since and recorded provided determined) found Johnston compiled that an status these An , occurred of tanker majority tires, Palmyra, which rodent-free, ballast decades. the caretakers Phoenix historically estimated supplies, nearshore intensive one anchors most can alien Islands. involved sponsored during along wreckage battle, sites bottles, that grounding served to during decimate information waters of Atoll list species during through are two fishing are and a the were WWII, Islands. and However and food, from inventory Baker previously 27 size During due and low and marine contained metal years as most noted towed-diver Kingman. Pacific can of a sites towed-diver chains, active exist regular a lying, vessels of to nuclear and the students and status native the arriving Atoll. was temporary also concerning the 6 racks, likely the to , debris Over at life, combined ship majority m focusing fuel. fuel in all military Biological surveyed the poorly probable concerted abandoned 2004 the fuel of wildlife by these survey migratory near testing PRIAs introduced. vessels. groundings Several 200 spill barrels, at either U.S. surveys from 12 Otherwise, majority surveys localized Palmyra season, forward of chartered, installations the remote m, on anthropogenic in marine and and Phoenix ef Howland, except in 2002 alien remnants alien Survey the forts marine was uninhabited rodent anchors steel the These shore by and monitoring of vegetation. is mid-1970s. Allied outbreaks always there track islands Comparable species and 1960s, not or the low marine ship is alien W pipes, military Islands, and eradication performed alien cryptogenic. provided birds, ake; Baker two , noted with PIFSC-CRED from DoD nets on Forces and was traffic species run lack and and objects remote NWRs. debris b) the documented species frames, many activities , their sea debris chains of have workers no 33 and Fuel during irregularly the All that is chemical southern beacons invasive in air data notable sites turtles, by limited guano ef risk Jarvis of in of noted T Areas atolls been spills base were 85% T forts able was The and and our are the the the the the for by at at of in page 345 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Table 12.1. List of marine alien species reported from the PRIAs since 2000. Source: Godwin and Vroom, unpublished data. SPECIES NATIVE RANGE STATUS IN PRIAs Sponge species 1 (Unidentified) Undetermined Cryptogenic, Palmyra Atoll only Sponge species 2 (Unidentified) Undetermined Cryptogenic, Palmyra Atoll only Areas Pennaria disticha (hydroid) W. Atlantic Alien Branchiomma nigromaculata (polychaete) Undetermined Cryptogenic, Johnston Atoll only Armandia intermedia (polychaete) Indo-west Pacific, Atlantic Cryptogenic, Johnston Atoll only Balanus amphitrite amphitrite (barnacle) SW Pacific, Indian Ocean Alien Bugula vectifera (bryozoan) Undetermined Cryptogenic, Johnston Atoll only Caulibugula dendrograpta (bryozoan) Undetermined Alien, Johnston Atoll only Didymozoum triseriale (bryozoan) Indo-Pacific Cryptogenic, Johnston Atoll only Halysis diaphana (bryozoan) Undetermined Cryptogenic, Johnston Atoll only Ascidia sydneiensis (tunicate) Undetermined Alien, Johnston Atoll only Diplosoma listerianum (tunicate) Undetermined Alien, Palmyra and Johnston Atolls Microcosmus exasperatus (tunicate) Undetermined Alien, Johnston Atoll only

Marine alien species in these remote areas have just recently become an issue of interest, due to survey efforts in other parts of the tropical Pacific. Efforts should be focused on minimizing the likelihood of coral reef habitats being exposed to marine alien species through the spread of organisms already established in altered habitats and transport of new species from outside of the PRIAs. This can be achieved by management efforts Pacific Remote Island directed towards all activities that have the potential for acting as mechanisms of transport, especially visiting ships and airplanes as well as the shoes and clothing of visitors.

Security Training Activities Security training activities are not practiced at any of the NWRs in the PRIAs. Data are not available for Wake Atoll, which is currently an active military installation.

Offshore Oil and Gas Exploration Offshore oil and gas exploration has never occurred, is not being contemplated, nor would it currently be feasible in the PRIAs. All are small low-lying atolls and reef islands surrounded by deep oceanic waters. The possibility of any commercial oil and natural gas formations is highly unlikely.

Other Crown-of Thorns Starfish Counts of the crown-of-thorns starfish (COTS), Acanthaster planci, were conducted around the U.S. Line and Phoenix Islands in 2001, 2002, and 2004 in conjunction with 50-minute towed-diver habitat surveys (see towed-diver method in the ‘Benthic Habitat’ section).

All 2001 and 2002 tows at Palmyra were resurveyed in 2004, and four new tows were added. Surprisingly, COTS were not observed at Palmyra in 2004, even though the survey efforts increased and they were recorded in 2001 and 2002 (Table 12.2). Aggregations of COTS were recorded by the USFWS and The Nature Conservancy divers in mid-2001 and 2002 off the southwestern terrace, but these apparently disappeared by 2004. COTS have yet to be observed by towed-divers at Baker, Jarvis, and Howland Islands.

page 346 adult COTSmayhideunderneathplatecoralsandotherstructures. predation. estimating at COTS, towed-diver 2001 12.7). surveyed and was km. in divers 2002 sightings at km. Johnston, the Kingman were In 2004. T able 12.2. Kingman Palmyra Jarvis Island Baker Island Howland Island Johnston Kingman LOCA 2001 Kingman the 63.2 in The surveyed Source: PIFSC-CRED,unpublisheddata. recorded and and In 2004, surveyed In 21.5% Atoll TION 2004 Atoll Atoll COTS in 2002, with and km 160% T divers addition Reef 2002 habitat otal numberofCOTSrecordedduringtowed-diversurveysandthetotaldistance(km)surveyedin2001,2002, 2004 in Counts surveys 37.8 of Reef 2004. 19.9 recorded of at survey were with tows, from 27.4 (T surveyed recorded increased 26.6 all T Johnston reported able km to COTS/km OWED KM of where The recorded most 13.8 26.6 17.8 10.1 1 N/A 9.9 2001. resurveying COTS 1.3 km of recorded two km year 12.2). towed-diver habitat 5.3 2.9 noticeable of COTS/km of habitat additional 44% that Atoll recorded , 2001 from T (Figure COTS/ COTS/ habitat habitat COTS COTS owed- 13% from was 162 and For towed-diver all at COTS 140 N/A of 4 0 0 0 2002, and2004.Source:PIFSC-CRED,unpublisheddata. Figure the The StateofCoralReefEcosystemsthePacificRemoteIsland coral 12.7. surveys T habitat OWED KM Recorded 27.4 10.8 1 N/A 5.4 31 1.1 typically along 2002 COTS those underestimate per COTS 378 N/A 28 0 0 0 towed two tows kilometer abundance appeared T OWED KM (km) 37.8 41.7 19.9 20.8 63.2 17 at white Kingman since 2004 due juvenile Reef COTS 752 182 to 0 0 0 0 in COTS Areas 2001, and page 347 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

CORAL REEF ECOSYSTEMS—DATA GATHERING ACTIVITIES AND RESOURCE CONDITION

WATER QUALITY AND OCEANOGRAPHIC CONDITIONS

Areas The health, functioning, and biogeography of the coral reef ecosystems of the PRIAs are primarily controlled by the oceanographic conditions to which fish, algae, corals, and other invertebrates of the ecosystem are exposed. This broad and diverse biological community is heavily influenced by time-varying ocean currents, waves, temperature, salinity, turbidity, nutrients, and other measures of water quality and oceanographic conditions. As these conditions change, so do the health, distribution, and species diversity of each reef community. Table 12.3 provides a list of long-term oceanographic and water quality monitoring programs in place in the central equatorial Pacific. Figure 12.8 shows the locations of many of these monitoring sites.

Table 12.3. Oceanographic monitoring systems in the PRIAs. SYSTEM VARIABLES MONITORED DATES AGENCY Deepwater CTDs* at select temperature, salinity, dissolved oxygen, Feb. 1999 - present PIFSC-CRED locations near the islands and chlorophyll versus depth to a depth of 500 m Shallow-water CTD* - temperature, salinity, turbidity Feb. 2001 - present PIFSC-CRED multiple sites each island/atoll Coral Reef Early Warning Enhanced: temperature (1 m), salinity, Feb. 2002 - present PIFSC-CRED System (CREWS) Buoys - 1 wind, atmospheric pressure, ultraviolet Pacific Remote Island Enhanced – (Palmyra) radiation, photosynthetic active radiation Sea Surface Temperature Temperature at 0.5 m Feb. 2002 - present PIFSC-CRED (SST) Buoys – 4 (Johnston, Howland, Jarvis, Kingman) Subsurface Temperature Temperature at depths between 0.5 m Feb. 2002 - present PIFSC-CRED Recorders (STR) – 24 and 5 m Ocean Data Platforms (ODP) temperature, salinity, spectral Oct. 2002 - present PIFSC-CRED – 2 (Baker, Jarvis) directional wave motion, current profiles Wave and Tide Recorders spectral wave motion, tides, July 2003 - present PIFSC-CRED (WTR) – 1 (Johnston) temperature Tide Gauges tidal fluctuations, sea level ? NOAA Ocean Service, Pacific Tides Branch Wave Monitoring Buoys – SE wave height & period, wind speed 1997 - present NOAA National Weather of Kiritami (Christmas) Island & direction, atmospheric pressure, Service, National Data Buoy temperature Center Satellite Remote Sensing sea surface temperature, winds, sea SST -1981 NOAA Satellites and surface height, ocean color SSH – 1992 Information, Hawaii Coastwatch Wind – 1995 Ocean Color - 1994 Model Fields waves / surface circulation NOAA National Weather Service, Wave Watch 3 Naval Research Laborator, Navy Coastal Ocean Model * CTD: Conductivity, Temperature, and Depth

page 348 Figure 12.8. oioiglctosi h RA. Map: A. Shapiro.Source:PIFSC-CRED. Monitoring locationsinthePRIAs. The StateofCoralReefEcosystemsthePacificRemoteIsland Areas page 349 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Ocean currents transport and distribute larvae among and between different atolls and islands, as well as define SSTs and available nutrient regimes. The equatorial region is home to several well-defined, though variable, currents: the NEC, ECC, EUC, and SEC. These currents are highly influential to the ecosystems of these islands and atolls: the ECC is the major pathway of larval transport and connectivity to regions of high

Areas biodiversity to the west; the deeper flowing EUC brings colder water and nutrient enrichment to islands nearest the equator (Figure 12.9); and the NEC and SEC are generally well mixed and nutrient poor. The magnitude and mean position of these current systems are highly dependent on El Niño/Southern Oscillation (ENSO), and ENSO-related changes to oceanographic conditions in this region are generally much greater than annual changes. Figure 12.10 shows this dependence; SST at all central Pacific PRIAs exhibit greater interannual variability and dependence on ENSO than seasonal changes, with the exception of Johnston Atoll which exhibits a more annual cycle similar to that experienced by the Hawaiian Archipelago.

Starting in 1999, hydrographic measurements and oceanographic parameters near these island and atoll environments have been

Pacific Remote Island measured using shipboard acoustic Doppler current profilers (ADCP), deep and shallow water conductivity, temperature, and depth device (CTDs), and an array of instrument Figure 12.9. Upwelling of cooler, nutrient rich waters originating from the Equato- moorings aboard the NOAA ships rial Undercurrent near Jarvis Island. Upwelled waters influence fish assemblages Townsend Cromwell (1999, 2000, and and distributions, coral growth rates, and a number of other components of the local 2001) and Oscar Elton Sette (2002 coral reef ecosystem. Source: Gove, 2005. and 2004) during regular scientific cruises for the PIFSC-CRED. Based on data from these cruises, as well as other satellite derived and in situ data sources listed in Table 12.4, a number of physical/biological linkages have been observed:

• A link between areas of upwelling EUC water with benthic habitat composition and distribution of reef fishes, especially planktivores, at Jarvis and Baker islands;

• Extreme interannual variability of water temperatures, upwelling, and other mixing phenomena (connected with nutrient supply and primary productivity) at Jarvis and Baker and Figure 12.10. Relationship of NOAA Pathfinder derived SST with ENSO Multivariate the dependence of this variability on Index (MEI) at the PRIAs of the central Pacific. Note Jarvis extreme dependence on the ENSO cycle; ENSO contrasting with Johnston’s annual cycle. Source: PIFSC-CRED, unpublished data. • Evidence of weak but consistent upwelling along Palmyra Atoll’s southern side and its connection to high biomass measured with bioacoustics; and page 350 (T in 2004, were Rare) Abundant, lines areas their inventorying REA and coral 78 2000 transects and by REAs Rapid Ecological Methods three oftheislands/atolls. per averaging Kingman Phoenix biodiversity Several CORALS BENTHIC HABIT ever-changing physicalenvironment. between to ecological of Continued in watertemperature(Figure12.1 and Palmyra • able the sites algal, site, establish oceanographic Line fish Poor associated populations relative using protocols and system. U.S. initiated and 12.4). are techniques surrounding specialists photo-quadrat/video Islands. Islands Lagoon were ecosystem Reef); (Figure about 2002, coral, assessments , 29 Phoenix water monitoring the concurrently Common, distribution, species abundance Sites sites quantitative Assessment conducted at focused DACOR large 2 and (Howland Assessments and 98 in During other 12.12). km Johnston A circulation along were have were the and TS REA conditions health American and backreef in diel the and U.S. invertebrate, are length, Line these Occasional, selected abundance, been on resurveyed (Dominant, three in surveys conducted estimating to variations and Between activities required linkages Phoenix analysis and transect broader Atoll surveys visually Islands assess years, within areas used 25 Samoa Baker); rapid and 1). the so m at in since at ecological population variation right lagic, the sametransect linesatKingmanReef. Figure 12.12. of coralbleaching. in Figure (Rose U.S. permanently a 2000 Palmyra Degrees °C The StateofCoralReefEcosystemsthePacificRemoteIsland indicates NOAA Line 12.1 in Atoll). assessments by water 1. structure, pathfinder backreef Islands position W A PIFSC-CRED USFWScoral biologist andtwoNOAA ater marked temperatures These Source: temperatures of (commonly (Jarvis SST the and 50-100 techniques PIFSC-CRED, unpublisheddata. . (REA) STR. Black condition indicate Date Island, and know recorded Highly each arrow m USFWS transects, poor as elevated Johnston include Photo: J.Kenyon. on covering in the by circulation overview the in Coral to situ daytime towed-diver PRIAs and assess Gardens) Atoll, between STR fishbiologists survey along of and recruitment CTD temperatures and a Palmyra including potentially and temperature shallow as 1,000-5,000 compared surveys monitor water studies Atoll, greater and the track Areas spatial each coral to CTDs U.S. and risk pe m at at 2 - page 351 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

as to initiate monitoring in locations Table 12.4. Summary of coral assessment and monitoring activities conducted by for which previous qualitative and/or PIFSC-CRED and USFWS in the PRIAs from 2000-2004. REA = rapid ecological as- quantitative data were available. In sessment, TDS = towed-diver survey. Source: PIFSC-CRED, unpublished data. 2004, protocols were revised to shift 2000 2001 2002 2004

Areas towards in situ quantitative coral LOCATION REA TDS REA TDS REA TDS REA TDS population data collection. At each Johnston Atoll NA NA NA NA NA NA 12 27 site, two of the three 25 m transect Howland Island 5 8 7 6 6 4 3 9 lines were videotaped for use in Baker Island 6 6 6 6 6 2 3 8 analyzing percent cover of primary Jarvis Island 6 9 4 4 5 4 5 11 benthic components and as a record of condition of the benthos. Each Palmyra Atoll 8 12 5 8 11 13 9 21 coral whose center fell within 1 m Kingman Reef 5 10 9 11 9 11 9 18 of the first two transect lines was identified to genus and assigned to one of seven size classes based on the estimated length (cm) of its longest diameter: <5, 6-10, 11-20, 21-40, 41-80, 81-160, and >160 cm. In addition, digital photographs were taken of coral species, including those within a broader area beyond the transect lines, to provide a more complete inventory of coral biodiversity at each site. The revised REA protocols are described in Maragos et al. (2004).

Towed-diver Surveys Broad-scale assessment of the shallow-water (<30 m) benthic habitats around the U.S. Phoenix and Line Islands were conducted during 2000-2002 (110 surveys) and 2004 (67 surveys) (Table 12.4) by the PIFSC-

Pacific Remote Island CRED. Survey paths in 2004 attempted to replicate selected paths covered in other years to monitor for change, or to fill in gaps in areas not previously surveyed. Benthic towed-diver surveys were initiated at Johnston Atoll in 2004 (Table 12.4). During each survey, two divers maneuver separate boards equipped with a digital video or still camera and temperature and depth recorders, while being towed behind a small boat. The tow path is recorded by a global positioning system (GPS) receiver onboard the boat, and a layback model is applied to the data to more accurately map the position of the imagery. Percent cover of salient benthic categories is quantified by whole-image analysis of still frames sampled at 30-second intervals. Towed-diver surveys bridge a gap between large-scale mapping efforts using satellite data and small-scale traditional belt- transects or roving diver assessments, thereby providing a mesoscale spatial assessment of reef habitats.

Permanent Transects At each permanent site, a 50 m surveyor’s tape was laid along each transect alignment marked with stainless steel stakes installed at 5 m intervals. A 1 m2 quadrat was laid sequentially and photographed along the entire transect at 1 m intervals for a total coverage of 50 m2 per transect. These data will be later analyzed for the same parameters as the REA coral census data including percent coral cover, size class distribution, frequency, mean diameter, and generic diversity. More detailed information on the choice and installation of permanent transects is reported in the NWHI chapter of this report.

Recruitment Plates In 2002, recruitment plates were attached to the base of Coral Reef Early Warning System moorings in the Palmyra Atoll and Kingman Reef (Figure 12.13) lagoons and to Figure 12.13. Divers install recruitment plates in the lagoon at Kingman Reef. Photo: K. Wong. page 352 bold type. in more in U.S. transects cnidarians species). species), (49 Howland Islands for 2000–2002, Following surveys Coral communitystructurefromREA Results andDiscussion sampled areas. and larval of Biennial data of moorings association larval Their assess to recruitment Island Baker at a these subsurface T each able biological temporal the Line species), than collected recruitment included: deployment collections plates location 12.5. (91 U.S. Islands and at 10% by Relative surveys will occurring with the sites species), dif calcareous ocean Kingman will data of Palmyra by Phoenix ferences Genera Baker coral in are colony enable as the address and surveyed the 2004 abundances well conducted with data highlighted in instrumented (80 fauna Jarvis deployments within instruments. abundance organisms. is comprising among Atoll Reef and as species), reflected levels coupling physical physical platform spatial in Island tallied (155 (170 belt- Line the the of of in in listed. All 2004. are location one least at in tallied was colony one least at which for genera in cnidarian USFWS and PIFSC-CRED by conducted sites 23 at surveys REA on Table 12.5. Area surveyed (m counted T T T Stylophora Stylocoeniella Stylaster/Distichopora Sinularia/Lobophytum Scapophyllia Sarcophyton Sandalolitha Rhodactis Psammocora Porites Pocillopora Platygyra Pavona Palythoa Montipora Montastraea Millepora Merulina Lobophyllia/Symphyllia Leptoseris/Pachyseris Leptastrea Hydnophora Heteractis/Stichodactyla Herpolitha Halomitra Goniastrea Gardineroseris Fungia Favites Favia Echinophyllia Dendronephthya Coscinaraea Chryptodendrum Cladiella Astreopora Alveopora Acropora GENERA otal cnidarians urbinaria ubastraea The StateofCoralReefEcosystemsthePacificRemoteIsland

Relative abundance of cnidarian colonies in the U.S. Line Islands, based

2 )

ISLAND JAR 0.00% 0.00% 0.00% 2.70% 0.90% 0.30% 0.00% 0.00% 0.10% 0.00% 0.00% 0.00% 0.00% 0.10% 0.00% 0.80% 0.00% 0.00% 0.70% 9.90% 0.00% 0.00% 0.00% 0.00% 0.80% 1.00% 50.4% 0.00% 9.90% 0.00% 20.0% 0.00% 1.50% 0.00% 0.00% 0.80% 0.00% 0.00% 0.00% 0.00% 1061 600 PERCENT OFCNIDARIANF VIS P ALMYRA A 0.10% 0.20% 0.00% 5.10% 1.90% 5.60% 0.10% 0.40% 0.00% 0.10% 0.00% 0.40% 0.00% 3.60% 3.30% 0.00% 0.40% 0.00% 0.80% 6.10% 0.00% 5.90% 0.10% 0.00% 0.40% 16.9% 24.4% 1.40% 1.80% 4.70% 1.00% 0.10% 0.00% 1.20% 0.60% 0.20% 2.00% 0.10% 0.20% 1 3684 T 1.0% 750 OLL AUNA KINGMAN A 1.40% 0.00% 0.10% 0.00% 0.00% 5.80% 0.10% 3.20% 0.00% 0.10% 0.50% 22.7% 4.40% 0.40% 1.10% 0.10% 3.70% 0.80% 0.00% 0.10% 0.10% 0.10% 0.40% 0.40% 0.20% 0.40% 0.00% 0.10% 0.20% 41.9% 1.60% 6.70% 0.40% 0.00% 0.00% 0.00% 1.30% 0.80% 0.00% 1.10% 5896 T 725 OLL Areas page 353 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Size class distributions of cnidarians for three of these locations in American Samoa are shown in Figure 12.14. Analyses of coral colony size-

Areas frequency distributions can reveal important characteristics of reef populations, and can be used as a tool to estimate the response of coral populations to the reef environment. These data will serve as a baseline to which future size class determinations at the same sites can be compared.

The total number of cnidarian corals and anemone species reported historically at the PRIAs and adjacent atolls are shown in Figure 12.15. The pattern clearly shows that coral and anemone species diversity is higher at Palmyra and Kingman, both in the path of the ECC. The ECC travels through much of the higher marine

Pacific Remote Island biodiversity region of the western Pacific and may be transporting larvae of additional species to the two reefs. The ECC and the two reefs may serve as important pathways and stepping stones for species dispersal to the atolls and reef islands of the central Figure 12.14. Size class distributions of cnidarians surveyed in REA belt-transects in the U.S. Line Islands during 2004. Source: PIFSC-CRED, unpublished data. Pacific. The total number of cnidarian corals and anemone species reported at the PRIAs and adjacent atolls are shown in Figure 12.15.

Figure 12.15. Coral and anemone species totals reported at all surveyed central Pacific reefs. Blue arrow shows the direction of the equatorial countercurrent (ECC). Sources: Maragos, 1974, 1995, 1997, 2004; Maragos and Jokiel, 1978, 1986; Mara- gos et al., 2003; Coles et al., 2000; J. Kenyon and S. Godwin, pers. comm. page 354 between 2000-2004. Montipora, documents period structure to early resurveyed and Jarvis, older the at Kingman, sites Rose; nine and the in transects Forty from 1999to2004 Shift incoralcommunitystructure concomitant the Ailinginae track Marshall Marshall 2004. each 2004, PRIA sites Palmyra PRIAs, permanent at five Kingman, changes specific were and over at and up sites the sites each providing These Atoll Baker Islands Islands. American increase Swains; to gradually decline a Pocillopora have three at sites. in at in at 50 two , Johnston, transects Baker coral Howland, the Palmyra; between Johnston an m times and to Many Figure of Samoa, now Republic established of opportunity , community or Acropora five Howland, two and through include 100 Porites Jarvis, of Rose, 12.16 three been 1999 sites year and and the the m of , hfe fo sal n nmru t lre oois Suc: aao ad Veit, and Maragos Source: colonies. large 2004. to numerous and small from shifted Acropora 12.16. Figure The StateofCoralReefEcosystemsthePacificRemoteIsland , Montipora Coral size and abundance at Kingman Reef site 5P from 2000-2004. from 5P site Reef Kingman at abundance and size Coral and Pocillopora corals have declined, while declined, have corals Porites coral has coral Areas page 355 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Figure 12.17 provides representative examples of changes at two of the older monitoring sites, KIN-5P in the east shallow lagoon of Kingman Reef

Areas and BAK-5P at Baker Island, over the same time period. At the Kingman Reef site, the 2002 photoquadrats along the permanent transect showed numerous COTS (Acanthaster) feeding scars and two of the starfish actively feeding on corals. The photos and other observations show that many types of corals are potential prey for the starfish at Kingman but that incidence of predation was higher on Pocillopora, Montipora and Acropora than on Porites when all four groups are present (as was the case at site KIN-5P). The data show a gradual shift to larger more abundant Porites over the four years of monitoring, with COTS predation on the other

Pacific Remote Island coral groups likely contributing to this trend.

These examples represent the preliminary findings at just two of 40 permanent monitoring sites in the PRIAs and neighboring reefs. The findings are also consistent with those of Timmers (this report) that document the movement of COTS towards the eastern lagoon of Kingman Reef in 2002, based on towed-diver surveys at Kingman Reef, and illustrate Figure 12.17. Summary of trends in mean coral diameter, mean coral frequency, generic diversity, coral percent cover, and Tridacna clam frequency at Kingman Reef the value of multiple strategies in site 5P (upper panel) and at Baker site 5P (lower panel) between 2000-2004. Source: assessing and monitoring coral reef Maragos and Veit, 2004. life at remote sites where dive time and dive days per locale are extremely limited. Monitoring corals and conspicuous macroinvertebrates at permanently marked sites has the advantage of isolating temporal trends in a set area. Analyses of all permanent monitoring data will allow comparison to coral data collected at adjacent REA sites at each of the PRIAs and help elucidate larger-scale geographic and temporal trends for coral population dynamics in the PRIAs. The findings should help NWR managers focus on assessing and eliminating anthropogenic stressors at these vulnerable reefs.

page 356 helped determinetherelative abundanceofalgalgenera(T over the Algal and availableforfutureanalyses. the studies for conducted. cover for endemic From PIFSC-CRED officeinHonolulu. on ameliorate U.S. species 1968). often Historically Results andDiscussion coralline algaeandmoderateabundanceofmacroalgaesuggeststhatthesereefsystemsarevery healthy the expeditions, Because of epiphytes. • allows phycologiststomoveof unable large • multiple diverswithvaryinglevelsofexpertisesubjectivelydeterminingpercentcoverinthefi • include: photoquadrats, the here only visits Quantitative Methods ALGAE Specimens Fish One in common percent US lack U.S. ability having years equatorial were monitoring amounts to biased situ an using transects person equatorial to conducted of of each Line species South algal no sampling sample written , to islands obvious or this cover one algal algal a it species quantitative Detailed collected towards decades sample and site analyzes is protocol perspective, problem, of islands and et phycologist in are impossible new collections by sampling algae islands. determination Phoenix (40-50 those the relatively al. concurrently algal of sample usually phycologists in to on (2001) U.S. large, photoquadrat if from algal undoubtedly devised the science. all growing environmental PIFSC-CRED blooms areas algal and minutes/year), Baseline equatorial performed images Islands PRIAs the per each to laid species close diversity from macroscopic added slightly f themaintransectintoshallowerwater;and sampling tell because US with benthic specifically of at or in Expeditions photoquadrat with the to how for a will algae for equatorial alien shallower underrepresents assessment algal . without considerable analysis the constant islands U.S. by away the first-hand analytical expeditions Most enable the or team, has U.S. species of the the and anthropogenic sampling first species Line present The StateofCoralReefEcosystemsthePacificRemoteIsland sampling for from PIFSC-CRED the data is ever inability during Phoenix water combined and invertebrates islands depth. long-term time. still remote will and experience aid data consistency on have the occurred the detail in to (Dawson state allow will Phoenix of a 2001-2004 the its Additionally the American of constraints contain for lack fish Previous short Islands; trained infancy aid tropical not phycologists reefs changes of to with data true all for of transect in able 12.6). in the our been on from at of distance easier , et Islands expertise the algal defining voucher combined many sets which . phycologists. the the the reefs however Samoa addressed reef al., knowledge phycologists Biannual , Pacific previous placed thoroughly quantitative occur U.S. species to diversity PRIAs species line, 1955; unique ecosystems were compares minimizes to away be algal in specimens over . Line , has work with established quickly on the the visits Preliminary before research Dawson, solely ecologically-based level. from of distributional habitats present determination, been the Advantages and the field. paucity analyzed have high away baseline microscopic to to past error benthic Phoenix record the established qualitative, (Preskitt the limited Oceanographic percent past The been and missions, from in 1959; and four fish of that PIFSC-CRED monitoring associated these and conditions. techniques assessments algal field percent by frustrated team. to are patterns the transect, years may Islands covers et T including turf four these are suda intermittent, ecosystems. sites notes eld; collections likely “fish al., and algal reveal have and stored factors: This results cover 2004) with are and of techniques throughout throughout give monitoring transects,” and presented to by questions However will research epiphyte crustose analysis focused in-hand method change of contain seeing having T divers at being using Areas allow rono, short were have algal from and the T . o , page 357 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Table 12.6. Algae of Johnston Atoll, Howland Island, Baker Island, Jarvis Island, Palmyra Atoll, and Kingman Atoll. Bold numbers indicate the number of photo-quadrats in which an alga occurred; italicized numbers indicate the alga’s relative abundance (rank) in relation to other algae occurring in the same photo-quadrat. Standard deviation of island averages are given in parentheses. Asterisks indicate algae found during the random swim that did not occur in photo-quadrats sampled. Source: Page and Preskitt, 2004. JOHNSTON HOWLAND BAKER JARVIS PALMYRA KINGMAN

Areas ATOLL ISLAND ISLAND ISLAND ATOLL REEF AVERAGE AVERAGE AVERAGE AVERAGE AVERAGE AVERAGE GREEN ALGAE Avrainvillea 8.33 (11.79) 11.46 (14.04) 9.26 (19.30) 4.17 (0.24) 4.68 (0.8) 4.88 (0.99) Bryopsis * 11.11 (16.39) * 3.9 (0.36) Caulerpa 12.50 (19.62) 6.25 (7.98) 3.13 (6.20) 3.70 (4.39) 3.15 (1.30) 5 (0) 5 (1.41) 4.25 (0.96) Codium 1.39 (3.4) 5 Dictyosphaeria 2.77 (4.10) 2.08 (4.17) 5.56 (6.8) 5.56 (8.61) 26.04 (23.75) 9.26 (12.11) 4.75 (1.26) 5 4 (1) 4.75 (0.35) 4.05 (1.03) 5 (0.41) Halimeda 10.42 (17.81) 33.33 (19.25) 26.39 (38.16) 27.78 (12.55) 70.83 (26.35) 71.3 (28.29) 3.08 (0.44) 3.21 (0.57) 3.17 (0.58) 3.65 (0.38) 2.82 (0.61) 2.72 (0.47) Microdictyon 34.26 (28.7) 4.01 (0.68) Neomeris 1.04 (2.95) 29.63 (22.86) 4 4.38 (0.59) Valonia 5.56 (13.61) 3.13 (4.31) 5.56 (8.33) 3.5 4.67 (0.58) 4.33 (0.58) Pacific Remote Island Ventricaria 4.17 (9.73) 0.93 (2.78) 2.67 (1.15) 7 RED ALGAE Amphiroa Chrysymenia 0.69 (2.41) 5 Dasya * Galaxaura 4.17 (10.21) 12.5 (35.36) 4.67 3.58 Haloplegma * Halymenia Hypnea Jania 1.39 (3.4) 3.7 (8.45) 3 5.83 (0.24) Laurencia 16. 67 (16.67) 3.63 (1.03) Peyssonnelia 4.17 (6.97) 2.78 (4.3) 5.21 (6.2) 11.11 (16.67) 3.75 (1.77) 3 (1.41) 3.75 (0.96) 3.83 (0.82) Wrangelia 37.5 (19.84) 3.73 (0.36) branched coralline 11.81 (23.42) 5.56 (10.09) 12.5 (23.42) 1.04 (2.95) 16.67 (23.94) 2.17 (0.58) 2.67 (0.94) 3.32 (0.25) 3 3.33 (0.84) crustose coralline 48.61 (29.69) 97.92 (4.17) 81.94 (20.01) 91.67 (7.45) 93.75 (9.71) 83.33 (20.83) 2.25 (0.50) 1.54 (0.32) 1.97 (0.79) 1.87 (0.47) 1.7 (0.24) 2.15 (0.35) BROWN ALGAE Dictyota 2.27 (7.54) * 36.11 (33.61) 5.56 (13.61) * 2.67 (1.89) 3.15 (0.6) 4.25 Lobophora 11.81 (20.56) 64.58 (28.36) 66.67 (39.09) 69.44 (25.09) 3.08 (0.67) 2.62 (0.54) 2.24 (0.99) 2.86 (0.89) orange crust 65.63 (34.05) 13.89 (18.63) 2.99 (0.63) 4.02 (0.99) Turbinaria * CYANOPHYTES 13.19 (18.96) 6.25 (7.98) 23.61 (26.57) 5.21 (8.84) 10.19 (10.02) 2.53 (0.98) 3.25 (0.35) 2.6 (0.25) 4.33 (0.58) 3.94 (0.14) TURF 92.36 (6.61) 93.75 (4.17) 79.17 (15.59) 94.44 (13.61) 94.79 (4.31) 97.22 (8.33) 1.21 (0.46) 1.87 (0.21) 2.18 (0.45) 1.51 (0.34) 2.31 (0.57) 1.32 (0.43) page 358 A Belt T more completereef-and archipelago-specifi belt-transect the them A REA structure andbiomassdensitiesbytaxa. atoll, method scattering wide-ranging mobile fishes; species of Several Methods associations. biological scales, biomass monitoring U.S. Inventories lagoon patchreefs,bays,backreefs,andshallowoceanicbankswherepresent. assemblages. monitoring prerequisite 2000 Reef) Field diversity byarea;andassessingef (e.g., target, baseline contribute fish Pacific Quantitative FISH used inmanyotherjurisdictionsacrossthePacifi rine the V ASSOCIA irtually pair pair diurnally USFWS current and resources. Protocol Pacific or surveys to ransect Protocol were overfishing, of and of Islands indicator 3) fish and the reef. presence complementary was all Atmospheric scuba densities to diver-observers data layer Stationary to Phoenix TED BIOLOGICAL has surveys monitoring was active largely lowest and species; statistical measure Islands and for assessment or replicated the fishes were Fish Habitat as diver-observers begun. will SPC, echo-sounds comprehensive too assessments Fish or PIFSC-CRED, scientific an at shallow-water habitat to recognizable length-class conducted qualitative aid are keystone (Baker strong (except over a be 4) integral dive Research point are MP correlations and types station Ongoing in , planned at T calculated, non-invasive owed-diver/video A a and conducted the damage, understanding time , sites assesement to basis ef counts broad-spatial Howland) surveyed W (SSL) fectiveness; part conduct species; primary or monitoring in ake with of permitting. conducted was (JIMAR). COMMUNITIES within working taxon. for assessment 2000, reef/bank; analysis reef shallow essential of to fectiveness oftemporalmonitoringmanagedareas. Island (SPC) the fish to sedimentation, each PIFSC-CRED’ estimated be an fish included identify sustainable underwater Islands, transects. and/or 2001, assessing activities primary assemblage This determined. arbitrary in patterns and of of scale, geographic assemblages. reef Their monitoring to parallel The collaboration surveys The StateofCoralReefEcosystemsthePacificRemoteIsland 2) for c fishspeciesinventories ateachisland,atoll,orreef. this shallow method 2002, quantify of patterns Farallon among Belt fishes mainly sound and for REA protocols focus growing in reef conducted c. swim The fish swims surveys all of transects living conjunction s prey and in (TDVS) structure reef data mission was fish fish have quantifi sub-region Further the size-frequency outer management. relatively REA 2004 on of community de at 2004 with size database resource migration Survey along fish involve typically complement distribution assemblages various site Mendinilla each were now protocol reef in at to to ed to the in assemblages changes); analysis to the selection. Johnston quantify quantify with three improve the slopes fish site, larger been be used types to various University PRIAs habitat used on response will in described U.S. document towed-diver/habitat to was [CNMI] 25 recording pelagic assemblages; and Related Pacific completed to enable of the provide around (>25 included: relatively at relatively m in the assessing Line are enumerate Atoll methods also PIFSC-CRED’ types An deeper abundance other long each were of scientific accomplished cm which to communities (Jarvis, coral initial species-specifi at used (Figure temporal Hawaii’ most objectives an fish transect possible present visual various total area. 1) , small-bodied conducted by large-bodied and estimate time-limited are reefs, species following REA species presence islands, the assessing the Palmyra understanding s length 12.8). as are protocols restricted Joint s Subsequent variability video; lines, diverse spatial around to PIFSC-CRED ecosystem oceanographic include: well and using similar by document of composition but inventory around c Institute [TL]) completion Fish scientists and recording Atoll, numerical survey numerical as sites and and (>50 and also the sonar components each to areas), to ecosystem in creating identifying and data assemble abundant 5) Kingman status temporal or methods included the reef of cm biennial impacts for . results island, simple Sound was where these Areas Each more at size- from from U.S. TL), size Ma of and and and and fish all of a a a - page 359 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

class specific counts of all fishes encountered (using TL) and identifying them to the lowest possible taxon, within visually estimated belt-transects: 4 m wide for fishes ≥20 cm TL (100 m2 area) on the initial swim-out, and 2 m wide for fishes <20 cm TL (50 m2 area) on the subsequent swim back. Transects lines were typically set at depths of 10-15 m. Reef ledges and holes were visually searched. Stations were completed on all sides

Areas of the island/atoll/reef, weather and sea conditions permitting.

SPC Protocol One SPC diver-observer conducted surveys in conjunction with, but at least 10 m away from, the two belt- transect divers. All fishes ≥25 cm TL that entered a 20 m diameter cylinder (area about 314 m2) during a 5- minute period were counted and identified to the lowest possible taxon. Individuals or groups were estimated to the nearest 5 cm TL size-class bin. Four replicate, 5-minute surveys were conducted at each station. Care was taken to avoid overcounting large transient or schooling species.

TDVS Protocol A pair of divers were towed about 60 m behind a small boat about 1 m above the bottom, at a speed of about 1.5 knots across about 2 km of coral reef habitat (during a 50-minute survey), generally at a constant isobath around the island. One diver recorded all fish ≥50 cm TL within a 10 m wide swath. The other diver quantified benthic habitat composition by type and macroinvertebrate densities. Fish were identified to lowest possible taxon and recorded in 25-50 cm TL size-class bins. Both towboards were equipped with digital cameras (video for fish and still for habitat) and temperature/depth/time recorders for more detailed future analysis. The towboat concurrently logged a GPS track position every 5 seconds. The fish diver performed a 360° scan during the first minute (plus another scan at the end of the tow), recording all large fish within visible range, and then completed 10 5-minute survey segments. Laboratory analyses of the digital videos recorded during Pacific Remote Island towed-diver fish surveys are ongoing.

Results and Discussion Preliminary results from the US Line and Phoenix Islands are presented here as full analysis of the data from these assessments is ongoing. The major effort by the PIFSC-CRED during its first few years focused primarily on field assessments.

U.S. Line and Phoenix Islands Results of the PIFSC-CRED assessments found that numerical densities of reef fishes were high at these islands. Small fish (particularly planktivores such as Pseudanthias, Lepidozygus and Luzonichthys) were quite abundant at Howland, Baker, and Jarvis Islands, particularly along the west side where upwellings occurred; fewer reef fishes occurred at Palmyra and Kingman. Distributional differences between islands were also discovered for these species (e.g., basslets). Densities of large fish (e.g., sharks, jacks, grouper, snapper, parrotfish) were higher in the U.S. Line Islands than in the U.S. Phoenix Islands or NWHI (Figure 12.18). Certain species of groupers, snappers and emperors occurred only at the southern or northern islands of the U.S. Line Islands.

Biogeographic patterns indicate that the majority of these reef fishes are widespread Indo-Pacific species that occurred at all five islands. A total of 480 fish species was recorded in the U.S. Line and Phoenix Islands in 2002. Total number of species was highest at Palmyra (343), followed by Figure 12.18. Density of large fish (>20 cm TL, all species pooled) in the U.S. Line and Phoenix Islands from belt-transects, compared to the NWHI in 2002. Source: Howland (302), Jarvis (252), Baker PIFSC-CRED (R. Schroeder), unpublished data. page 360 trimaculatus, densities in grouper (Carcharhinidae, snappers carnivorous on and diver Preliminary of thesefiveislands. to U.S. Phoenix Ocean occurrence central of that of characteristics but consistent genotypes Islands. found and U.S. enhanced and result densities phenomenon surveyed in Islands dispersal from an that finned includes three-spot one evaluated of one at also and Kingman (241), other the the this most otherwise the substantially centered trends 35% group oceanic the found. snappers more coral Line each of exceptional incongruent and fish PIFSC-CRED in Phoenix domino, Pacific group species U.S. nominal than the U.S. (Serranidae) the in the other at (where patterns. of Islands by The and with reef of Kingman of species work productivity the data remoteness Palmyra These fish. top within within of analysis recently reefs. the damselfi conditions in these U.S. French (Lutjanidae), in Phoenix Indo-Pacifi species and Phoenix fishes distribution geographic U.S. ecosystems of were French south-central U.S. Dascyllus in carnivores American that PIFSC-CRED. Jacks very on species, for Sphyrnidae), conservation is Islands higher the Line islands consistently likely The islands A Line four the dif (225). (where the recorded data Polynesian is sh, described collaborative with unique Pacific the few were of . ferentiation Polynesia Islands U.S. complex) define (Carangidae), Islands, needed. U.S. of and conducive c and systematics group many the families Within previous the to fishes separation include auripinnis, U.S. Samoa of numerical for Dascyllus may indicating Many and Line recorded most Line Phoenix Phoenix external a focused Islands mtDNA islands among towed- sharks Pacific genetic are in recent some value other jacks gold- adds be high Line This and species. was The (the and and that and new earlier the the a the study or of to in surveys a signifi a Phoenix revision records study larger only 2002, ily unpublished data. Figure sampling with from cant A The StateofCoralReefEcosystemsthePacificRemoteIsland area were few to one 2004) ecological 12.19. towed-diver Islands for component of further fish centered of conducted), the species and focused). Density co-occurrence specimens surgeonfi Johnston appear clarify surveys geneticists in presence of of Micronesia. common Inter-island taxonomic south-central to 78% Atoll sh conducted be were genus (2004 at at for at at large the Baker collected the these a only). relationships This in western-central dif fish Ctenochaetus junction University the Pacific ferences , (>50 islands 72% is Source: U.S. exemplifi (by cm of at Line endemic permit) TL, two were Jarvis, PIFSC- in of and and all species , California-Santa Cruz biogeographic ed which Pacific species Phoenix found: to CRED by species in 60% help the 2002 demonstrated patterns pooled) (S. species over at Islands contribution understand and Howland, not Holzwarth), 96% areas, by found Areas (2001, 2004 were and fam at - page 361 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

greatest densities, followed by sharks and then groupers (Figure 12.19). When the data were grouped by island, it was evident that the abundance of large predators fluctuated considerably between years and sites (Figure 12.20). In general, large fish densities appeared to increase at Howland and Baker from 2002 to 2004, while decreasing at Kingman and, to a lesser degree, Palmyra (for three of the four families). Areas Howland and Baker Islands (U.S. Phoenix Islands) Baker Island is characterized by a subsurface eastward flowing EUC that lies beneath the westward surface flowing SEC. This causes an upwelling of cooler, nutrient- and plankton-rich waters along the west side of the island, which contributes to enhanced ecosystem productivity. In 2002, the density of planktivorous fishes was higher along the west side (Figure 12.21).

Fish were resurveyed at Howland and Baker Islands from January 21-24, 2004. Observations from 11 belt- transects and SPCs (five at Howland and six at Baker) yielded the following preliminary results: 164 and 166 fish species were documented at Howland and Baker, respectively, compared to 126 and 127 species seen in 2002. Species richness averaged 118 and 117 species (median among stations) at the respective islands.

Together with observations from other dive teams (e.g., towboard fish team), a grand total of about 210 fish species were recorded at Howland and Baker. Both numerical and biomass densities of resident reef fish species remained Pacific Remote Island high (as observed on the 2001 and 2002 surveys). A trophically diverse fish fauna, representative of healthy coral reef ecosystems, contributed to the high biomass. The body size distributions of fishes spanned a broad range of sizes, further indicative of a healthy fish assemblage. Three species of small- bodied (<10 cm TL) zooplanktivores were numerically dominant, including two species of anthiine basslets (Pseudanthias bartlettorum and Luzonichthys whitleyi), and the basslet-like damselfish, Lepidozygus tapeinosoma. Major contributors to biomass included numerous medium- sized (10-20 cm TL) benthic fishes (both herbivorous surgeonfishes and parrotfishes as well as benthivorous carnivores) and diverse, large- bodied (generally 25-60 cm TL) piscivores. In terms of biomass, the mid-water piscivore community near the reef was dominated by twinspot snapper (Lutjanus bohar), grey reef sharks (Carcharhinus amblyrhynchos), reef whitetip sharks (Triaenodon obesus), and several Figure 12.20. Density of common large fish (>50 cm TL, all species pooled) by fam- ily from towed-diver surveys conducted in the U.S. Line and Phoenix Islands (2001, carangids (primarily the black jack 2002, 2004) and Johnston Atoll (2004 only). Source: PIFSC-CRED (S. Holzwarth), [Caranx lugubris], rainbow runner unpublished data. [Elegatis bipinnulatus], and bluefin page 362 nigricans rubroviolaceus of Jarvis, including ( grouper years. from ( on 2004. especially Jarvis including that and Fish Jarvis Island(U.S.LineIslands) remained abundantattheU.S.PhoenixIslands(Figures12.19and12.20). and islands. sexfasciatus Sphyrnidae blacktip gray undocumented while Islands. Quantitative and pygmyangelfi densities Baker were In occurs particularly in substantially Howland The fish biomassnearreefs. albomarginata] slenderspine flagtail fasciatus argus the dozen trevally Density V Carcharinus ariola 2002, parrotfish the American peacock six three were stock reef 130-180 ], decreasing density found by Jarvis, west except species (e.g., blacktip Groupers Islands SPCs, of and grouper louti ( [ In ], Snapper ) andspotted bristletooth( two- C. and Cephalopholis PIFSC-RED the a sharks surveyed of surgeonfi in along 2004, abundances ). coral where signifi side ) surveys whitetip ) melampygus T were deep between grouper Baker black were epinepheline like amblyrhynchos T were to higher utuila, for of cm ), all ), winspot Samoa shark grouper of in the grouper and four-fold at of the also cant ( were the Howland re-surveys [ and hind groupers shes. Carcharhinus TL C. substantial grouper observed; water found jack most also Islands the shes at terms west reef of towed-diver Guam, Pacific twinspot fining), per [ was than shark Jarvis urodeta Cephalopholis apex contributed Howland large very island. snapper [ [ ( urodeta Epinephelus C. appeared and somewhat of Caranx was sharks in at ]). side; dif site), 148 and the appeared large of (primarily groupers Baker abundant ferences predator miniata numerical Saipan). at fish [ steephead at Island of ) or Gracilla relatively most ], snapper fishing A CNMI, and U.S. mean in Baker in sites The simply but ), sites ( ( both amblyrhynchos were lugubris schools C. fish and and Lutjanus half 2002 2004 , coral to white Ctenochaetus marginatus while relatively from common ], common seemed Line melanopterus density established , team presence. completed is at natural densities ( low and fish/100 m side around Figure they hind Lutjanus parrot in March Islands. all ) none at tips bohar and line saw and The StateofCoralReefEcosystemsthePacificRemoteIsland (size 171 Howland, sites of were to ( were and unchanged 12.21. C. ( Baker population T large with ( the were one 26-27, primarily be Chlororus riaenodon of ), 2 (from ) appeared ). miniata by by bohar The surveyed This were quite circle also but and Source: bluefi slightly the Island the large Relative towed-divers the fish encountered and total 34 there could 2004. T ); found proportional EUC, bridled very large PIFSC-CRED riaenodon ), n remained represented (2002), some families) humphead fluctuations. from black and PIFSC-CRED (R.Schroeder),unpublisheddata. microrhinos trevally to obesus less number distribution in were reflect abundant resulting increase ). to (>50 Preliminary 2004. peacock parrotfish individuals the indicating common be jacks in to ), also in by ( obesus cm high abundant dif C. density 2001, of high were of wrasse The 2004. by ferences ( the ), in melampygus in coral in C. TL). at planktivorous The galapagos at higher grouper some whitecheek enhanced 2004 February levels both lugubris most observations ( along fish 2002, ), of Scarus common were all majority Planktivores and reef planktivores Jacks ( Chelinus on sites. densities REA islands. at of in common >70 the and recorded hammerhead the ( Howland fish illegal ) which C. frenatus fishes were of sharks were upwelling ). team. (typically west of Sharks, 2004 southeast surgeonfi cm argus 2001 species along Snappers undulatus at sharks from fishing Bigeye from exceeded very TL. species common that remained side at in ). and ( ), or Overall, the C. belt-transect six the station; Only mostly redlip common previous L and 2002, >10 documented sh observed western galapagensis sharks yretail pressure Baker than reef jacks belt-transects U.S. ) were ( were Acanthurus productivity at six at individuals 60 abundant, mean parrot terrace large indicated gray in Howland all Phoenix ( Islands, grouper upwelling species (Family Caranx cm at rare flagtail surveys earlier years, Areas sites, (e.g., were 8-161 both reef fish TL. ( S. at at in ), page 363 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

Towed-diver fish data from Jarvis Island suggest that densities of jacks were lower in 2004 compared to previous years, groupers and snappers were of similar range, and sharks were higher (Figure 12.19). Sharks were represented by gray reefs (Carcharhinus amblyrhnchos), blacktips (C. melanopterus), and whitetips (Triaenodon obesus). In addition, a school of 15 large (400-500 cm TL) great hammerhead sharks (Sphyrna

Areas mokarran) were encountered in deep water on the edge of the southeast terrace. A relatively high number of manta rays and a single whale shark (Rhincodon typus) were also seen in this area of the reef. Turtles were also abundant, with over 60 recorded during tow surveys in 2004. With the exception of jacks, large fish remained abundant at Jarvis in 2004.

Palmyra Atoll (U.S. Line Islands) Fish were surveyed around Palmyra Atoll from March 29 to April 1, 2004. No surveys were conducted in the heavily dredged, sediment-laden lagoon. Preliminary observations from 10 belt-transects and 10 SPCs, all re-surveys of sites established by PIFSC-CRED in February of 2001 or 2002, indicate that stock abundances appeared relatively unchanged from the high levels recorded in previous years. The total number of coral reef fish species documented at Palmyra by PIFSC-CRED was 193 in 2002 and 209 (from 34 families) in 2004. Large fish were common but not as abundant as at Jarvis. Sharks (mostly blacktip, followed by gray reef and white tip) were small (130-160 cm TL), and occasionally seen, but seemed to be slightly less common than two years earlier. About 16 species of grouper were seen at Palmyra, with most common being the peacock grouper (Cephalopholis argus; 15-35 cm TL) which were present at every site. The flagtail grouper (C. urodeta) and the camouflage grouper (Epinephelus polyphekadion) were somewhat less common. Jacks were common at all sites, including large giant trevally (Caranx ignobilis; about 100 cm TL), bluefin trevally (C. melampygus), and black jacks (C. lugubris). Snappers were very common at Palmyra, with the twinspot snapper (Lutjanus bohar), humpback snapper (L. gibbus), smalltooth jobfish (Aphareus furca), and onespot Pacific Remote Island snapper (L. monostigma) observed at every survey site. Most twinspot snapper ranged from 30-40 cm TL, smaller than those seen at Jarvis Island. Humpback snappers were observed in loose aggregations of up to 50 individuals.

Preliminary results from towed-diver fish surveys indicated differences in mean counts (numerical density) between fish families and years (Figure 12.20). Snappers were most abundant in 2001. Densities of jacks were least abundant in 2002. Shark abundance was fairly consistent between years, although a predominance of juvenile gray reef sharks (85% were less than 150 cm TL) may be indicative of fishing pressure. Most reef whitetip and blacktip sharks appeared to be mature size. Pooled grouper species at Palmyra varied in number among the three surveyed years. Most of the grouper >50 cm TL recorded were lyretail grouper (Variola louti), but relatively large individuals of the genera Epinephalus and Cephalopholis were also counted.

Kingman Reef (U.S. Line Islands) Fish were surveyed at Kingman Reef from April 2-4, 2004. Preliminary observations from nine belt-transects and nine SPCs, all re-surveys of sites established by PIFSC-CRED in February of 2001 or 2002, indicate that stock abundances appeared relatively unchanged from the high levels recorded in previous years. However, overall fish density for both small and large fish appeared to be lower than in previous years, with the possible exception of fish assemblages along the outer reef slopes. The total number of coral reef fish species documented at Kingman was 165 in 2002 and 187 (from 35 families) in 2004. Sharks were less abundant and smaller at Kingman than at either Jarvis or Palmyra. An average of only one or two sharks was seen at most sites (none at some sites), compared to approximately 12 sharks observed on a typical dive in 2002. The same three shark species–reef whitetip (most common), blacktip, and grey reef–were most abundant. Groupers were not as abundant at Kingman compared to Palmyra or Jarvis. The peacock grouper (C. argus) was most common, observed at every survey site. Also relatively common were the flagtail grouper (Cephalopholis urodeta) and the camouflage grouper (Epinephelus polyphekadion). Six species of jacks were observed at Kingman but were generally less abundant than at Palmyra or Jarvis. Most common were bluefin trevally (Caranx melampygus) and black jacks (C. lugubris). The twinspot snapper (Lutjanus bohar), observed at every survey site, was the only common large snapper at Kingman, although most were smaller than those observed at Palmyra or Jarvis. Humpback snapper (L. gibbus) and smalltooth jobfish (Aphareus furca) were present in lesser numbers. No maori wrasse (Cheilinus undulatus) or bumphead parrotfish (Bolbometopon muricatum) were observed by any dive team, consistent with previous years. Like Palmyra and Jarvis, Kingman is a NWR and levels of historical or recent fishing are unknown. There are, however, occasional unconfirmed page 364 low-intensity that because within the body thought coastlines, 300 SSLs Methods SSL amblyrhychos medium-sized and most of terrace, due large Hawaii areas operations generated the At populations. ( bluefi Standing densities (e.g., distributed was Preliminary Fish Johnston should helprevealcausativefactors. in monitoring disturbance chanos steephead cm sexfasciatus all Preliminary need toincreaseitsenforcementpresenceatKingman. 2004, reports C. safety large Johnston declined TL. survey ignobilis location Protocol in the a m low; were n condition snappers jacks are bigeye school the compared that part during ) than trevally of dearth areas to Gray fish were many considerations. of biomass Atoll (U.S.LineIslands) comprised of atoll fishing, sub-optimal play Indo-Pacifi period parrots surveys to were where in surveyed at results and ) the observations of ) Atoll, may to reef densities recruitments were of the emperor were general, ) wereseen,aswella2.5mtigershark( the reef Kingman, with also sharks, (1 survey abundant of but the (Figure ( an near about Caranx sharks 1 inaccessible to difficult juvenile have there day they resident towed-divers especially were ( sharks of also fewer uncommonly U.S. important encountered. seen, more Chlorurus planned earlier of the or at c , and sites 200 the several form and, 12.20). survey species taken pelagic was Monotaxis appeared total were conducted Line compared melampygus Johnston 2004 Hawaiian large survey common The as from life-stages, were that tow rise bigeye species survey within the for a together in at lack and role seen. place microrhinos tow large towed-diver to fish, conditions previous belt-transects very team occur Jack into communities Kingman the (abundant completed mesopelagic rare conditions observed. the Phoenix of in Grey team relatively lagoons. jacks years, Atoll species, to in grandoculis in received U.S. the northwest large appeared at and the data in was Solitary other previous at ) including 2004. the with Kingman were 2004, photic was reef years, a ( for Line coastal shark C. and lagoon able with Galapagos ), may for very in fish Islands. comprehensive 26 encountered dive The StateofCoralReefEcosystemsthePacificRemoteIsland were The including low sexfasciatus the Due able comparatively sharks The of Hawaii, infrequently boundary backreef and Islands large however Palmyra. surveys swell zone decreased surveys years to ; low and densities suggest older isolated , small observed first bluespine ecosystem teams. mostly Reef and more tow to to large SPCs , black jacks dive possibly a Johnston ( that at (Figure Carcharhinus young-of-year-sized the time will endemics, fish, one sharks over large between during night. and suggest abundant reef community fish possible atoll due The at complicated appeared there on ) closest document Fish jacks observed were visibility on were Galeocerdo cuvier shrimp, scarcity by a each more unicornfi PIFSC-CRED forereef northwest the data whitetip as 12.20). to lower variety inhabited ( the due the Carcharhinus SSLs had the in assemblages ( few that a forereef, mostly recorded. Caranx 2002 were neighbor of 2004. backreef). of from recent in major initial . . to PIFSC-CRED and almost of low densities amblyrhynchos 12 the Although 2004 snapper large-bodied long-term sh, In and are habitats, density of No the both the either and storm towed-diver Hawaii, stations at squid habitats by survey bluefi baseline The lugubris link Naso common large poaching often but surveying Johnston than general fish, islands) no data substantively jacks 2004. In , absent galapagensis between shallow Conclusions increased. not n ). of shark, at swell, that the shallow-water addition respectively schools (e.g., in unicornis time is boundary small trevally indicate sharks on Johnston throughout ) and extensively past assessment a large species. were occur from in The scarcity were activity reflection ) compared surveys the than surveys or jack the annual pelagic appeared backreef reef in years. zooplankton rare, of to and mostly ( swell PIFSC-CRED January coral exposed ). size; in Caranx that conspicuously and the self-seeding Parrotfish, bigeye community a sharks . are . ) are waters number Overall, Carangids and and of to large , The the It suggest were forereef, waters large grouper reef of and hampered Milkfish no reef therefore in to is and decadal) smaller more juvenile to the several atoll, gray jacks 2004. melampygus other very USFWS 12-16, giant likely fore-reef fish this fish be deeper restricted groupers ecosystem, submerged and infrequent, of and especially numerical similar SSLs a densities in reefs because ( covering is possible diversity reef found ( small biennial Chanos islands, than Caranx reflects like trevally shift absent During limited fishes. higher widely trends robust where diving 2004. along Areas than may and fish are the ( 50 C. or or to to to in ) page 365 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

trophic levels. The observed horizontal migration towards shore at night of the Hawaiian community also suggests SSLs may play a significant role in the coral reef ecosystem through input of biomass and nutrient cycling. Little is known about the character of boundary community SSLs associated with islands outside of the main Hawaiian Archipelago. To improve our understanding of these communities, island-associated SSLs

Areas were investigated using ship-based echo-sounders during a PIFSC-CRED cruise to Jarvis Island, Palmyra Atoll, and Kingman Reef in 2004.

Results and Discussion Jarvis Island Echo-sounder surveys were conducted along the southern and western shores of Jarvis Island during afternoon and night/morning hours on March 26 and 27, 2004. Survey lines were run approximately 400-600 m from shore in waters 300-600 m deep. Each line was conducted at approximately 00:00, 02:30, 05:00, and 14:00 hours.

The surveys revealed a clear diurnal trend in the occurrence of a layer of surface-associated biomass. At night, the layer is generally concentrated in the top 130 m, just above the thermocline and centered on the chlorophyll maximum observed during CTD casts (Figure 12.22). The layer’s nighttime density was more or less constant along both transect lines. During the day, the layer was absent from the southern transect, but was very dense towards the northern end of the western transect. Because trawling for samples was not feasible during this cruise, the biological composition of the layer is presently unknown.

Palmyra Atoll

Pacific Remote Island Echo-sounder surveys were conducted along six transect lines around Palmyra Atoll between March 30 and April 1, 2004. Transect line lengths ranged from 4 to 9 km. Two lines were run along the edge and on the southwestern bank of the atoll, one along the southern shore, one along the edge of the southeastern bank, one along the northeastern edge of the bank, and one along the northern shore. All but the northern and northeastern transects were run multiple times during the same and/or Figure 12.22. Echogram of the SSL at Jarvis Island. Depth is plotted on the y- axis, horizontal distance on the x-axis (2.6 km). Red regions represent the bottom. successive nights, as well as during Source: M. Lammers, unpublished data. the day.

The data revealed that a dense SSL exists around the entire perimeter of the atoll at night. This community resides primarily along the atoll’s steep slopes at a depth 120-180 m and is centered on the thermocline (Figure 12.23). A secondary, less dense layer of organisms occurs above the thermocline in patches. Opportunistic observations made during transits perpendicular to the atoll’s edges indicate that both layers extend well offshore (4-6 km) along the southwestern side of the atoll, but Figure 12.23. Echogram of the SSL at Palmyra Atoll. Depth is plotted on the y- that they are restricted to less than 2 axis, horizontal distance on the x-axis (2.6 km). Red regions represent the bottom. Source: M. Lammers, unpublished data. page 366 ( Mysticetes habitat knowntobeexploited bybeakedwhales. the tentatively Although pers. comm.)reporthavingseen both atoll. seen resident. Palmyra common ecosystems. operations marine V MARINE MAMMALS the lackofawell-defi EUC the terrestrial of contribute upwelling is It observed ateitherJarvisorKingman. than distribution The primary channelintoandoutoftheatoll,similiartothatseenatPalmyra. most mid-water eastern An collected opportunisticallyononeoccasionwhiletransitingalongthesouthwesterncornerofatoll. of perpendicular the Echo-sounder Kingman Reef made alongthissideoftheatoll. the and enriched km Megaptera ery is the the islets assessment of southwestern southwest. oceanographic atoll interesting echo-sounder the little T Kingman. on fshore prominent nor Anecdotal . top day truncatus mammal and prevailing successive and between species that by any is ecosystem from not Palmyra of . organisms. to identifi revealed of presently were nutrients novaeagliae along southern The a Kingman. the . Of terrestrial-based the also directly surveys to steep the The Coincidentally patch of accounts Furthermore, the odontocetesencountered,bottlenosedolphins ( sighted. surveys mid-water ed and mid-water note reported side perpendicular April the data tradewinds EUC the also supports and two days as that ned SSL known flowing pinnacle are P sides other was These where that observed, data . Cuvier supports Repeated were 2-4, collected parallel bathymetric electra and ) over likely all have were Populations from observed three only of sides. biomass productivity collected about sites 2004. is a out the patches conducted there. , ’ a the blow s rising Palmyra ecosystem. a also signifi several USFWS observed transects ever on three-day for transects beaked of locations a contributing lagoon’ T at This speciesisknowntofeedprimarilyonsquid. beaked visited sightings atoll. . truncatus the population A past the the a T from one conditions along tentative cant from occurring been wo indicates major were marine of lagoon. observations three visits. supports staf Both observed s whale 50 location along transect do the were at waters period seabird the are whales were the conducted The StateofCoralReefEcosystemsthePacificRemoteIsland very made f to A seabird Johnston Atoll. have familiar andspermwhales( northeast, sites factor night mammal of minimal explanation 100 quite ocean southwestern there. Similarly conducted The ( (upwelling, four that Ziphius melon-headed localized run and flow a during visited lines or there. cetacean reveal may population. and main behind distinct dense, a colony transect with more at of were floor Jarvis out. SSL in or populations , daytime melon were 19:00 cavirostris probably also consecutive USFWS these channel reduced Palmyra by that and . Palmyra, at for animals . consistently deep Finally is the oceanographically thermocline-associated It and populations 18:00, Nutrient corner the This lines run this be conspicuously each ranged is headed and productivity whales areas, transects Nitrogenous located Palmyra waters) PIFSC-CRED present Physeter macrocephalus staf , parallel observation into creating finding (J. influx ) occur Kingman in along at location 21:00, was on of T Maragos, the f days inputs in ursiops truncatus the 01:00. the familiar recent whales ( the located Peponocephala depth of found almost at U.S. associated found is appear the lagoon revealed to atoll. at 0:00, suggest a nutrients other Johnston, observed rather is from Reef net the missing some and southern PRIAs. visits is quite between Echo-sounder ( and with at near pers. Peponocephala at This cruises. and at that movement atoll’ hand, lies to these Kingman is Jarvis organic signifi ecologically the that only Jarvis be distinct during PRIA community influenced the at once the with obs.) again on along there, s Howland, Although much equator ) Kingman 80 these slopes two is and southwest electra dispersed the cant were by southwestern are local Island a Humpback towards m sites. report ) therepreviously outflows coincided Reef. the in most sources southwest particularly forested of eastern more and characteristic as data populations the . water ). and PIFSC-CRED island by no Baker and electra that Jarvis there (M. may having far 130 These parts density In A neither systematic productive patches the side were two is from probably the addition, Rauzon, skeleton sides was network towards with , m. explain whales or middle Jarvis, are corner side ) fed Island of of along Areas were most were seen were atoll also The and are the the the the not the no by of of of is . page 367 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

modern records of this endangered species there. According to observer accounts, all social roles typically found on winter mating grounds are present at Johnston, including mothers and calves with male escorts, competitive groups, and singers. It is speculative, but Johnston Atoll may represent a relatively newly established or re-established mating ground of the north Pacific humpback whale population. Areas The role that marine mammals play in the ecosystems of PRIA sites is presently not well understood. The high metabolic needs characteristic of mammals suggest that their role is probably not trivial. Recent modeling work on the impacts of great whales (including sperm whales) on global primary productivity has revealed that up to 62% of the ocean’s primary production may have gone to sustaining whales during pre-whaling times (D. Croll, unpublished data). Although similar estimates are not presently available for smaller cetaceans, this does suggest that a more detailed assessment of the composition and abundance of marine mammals in the PRIAs is warranted.

MARINE INVERTEBRATES OTHER THAN CORALS Historically, biodiversity assessments and monitoring programs for coral reefs have focused on cnidarians and fish, so less information is available for other invertebrates. However, the sponge, mollusk, echinoderm, crustacean, annelid, bryozoan, and tunicate fauna associated with coral reefs are also integral components to the overall biodiversity.

Most data collection efforts in the PRIAs have focused on fish, ciguatera, algae, and corals, but some information on non-coral invertebrates is available from baseline faunal surveys conducted in the mid-1960s (Brock et

Pacific Remote Island al., 1966; Amerson and Shelton, 1976). A presentation of the species richness of non-coral invertebrates for the PRIAs will be a compilation of studies conducted since the early 1900s and collections held at the Bishop Museum. The information reported here will only provide data Table 12.7. Species richness data for selected non-coral invertebrates. Sources: Godwin, 2004, 2002; Coles et al., 2000; Brock, 1979; Amerson and Shelton, 1976; for porifera, hydroids (Cnidaria; Order Brock et al., 1966; Kay, unpublished data. Hydroida), anemones (Cnidaria; Order PHYLUM PORIFERA 20 Actiniaria), selected mollusks, and echinoderms. To date, the data for all PHYLUM CNIDARIA invertebrate groups are preliminary Order Hydroida 5 but these particular species have Order Actiniaria 9 the most complete species richness PHYLUM MOLLUSCA data. Only species recorded from Class Gastropoda 247 coral reef and associated shoreline Class Bivalvia 47 areas will be included in the species PHYLUM ECHINODERMATA 67 richness data provided in this section in Table 12.7.

The PRIAs possess marine invertebrate diversity and richness contained in habitats with a broad spectrum of anthropogenic influence. Locations such as Kingman Reef in the northern U.S. Line Islands represent locations with very little anthropogenic influence, while the coral reef habitats at Johnston Atoll are heavily impacted by alterations. Despite this vast difference, each of the PRIAs provides a refuge for marine invertebrate communities that recruit from a variety of source locations. For example, the giant clams Tridacna maxima and Tridacna squamosa can be found commonly in most of the PRIAs, with Kingman Reef possessing the most abundant communities. These clams have been heavily depleted in other parts of their range and the PRIAs represent a refuge for these species, if existing management and protection remains in place there.

page 368 its refugesagainstcommercialfi be will continueto and of initiative, anthropogenic Hawaii. reefs The CURRENT CONSER • • • • • • • • • summary statementscanbemade: Johnston complementary (including NOAA, Overall ConditionsandSummaryof vulnerable tounauthorizedfi Although stress, alienspecies,andperhapscoraldiseasetoresidentreefecosystems. Abandoned sharks, jacks,groupers)fromearlieryears. Surveys levels ofcoralspeciesdiversityatPalmyraandKingman. Coral Baker High elsewhere inthePRIAs. Substantial There ispresentlynoknownharvestingforthecoralandlivereeffi apex predatorscommon. negligible Levels appear toremainquitehealthyandproductive. Based its adjusted discourage USFWS NWRs that USFWS, densities , Howland)wereassociatedwithupwellingscausedbyimpingementoftheEUC. The larvae the on of Atoll, many are conducted unauthorized uninhabited accordingly to beginning the USFWS USFWS and WWII COTS light. co-managed stress transported be managedasno-takeMP and unauthorized new fish underwater Bishop of NOAA material, assemblage small Rose Overall, records) will and to predation at in V Museum, these to A Kingman atolls prepare 2004. NOAA fishing will TION MANAGEMENT planktivorous protect Atoll. in in survey reef military and access ECC continue ecosystems, shing andcollectingduetothelackofon-sitesurveillanceenforcement. the and The shing, destructivefi on will Based around and composition fish and assessments in resident U.S. from corals continue islands techniques USFWS 2004 construction, and coordinate assemblages other Analytical Results to on Line the fishes the poaching suggest collaborate As, has fish The StateofCoralReefEcosystemsthePacificRemoteIsland scientists and the serve U.S. western to will Islands, surveyed and catch-and-release and monitor and found been inventories, at work the work Line occupation, as an most ACTIVITIES shing, andunauthorizedfi of at seabird initiated development and Pacific with observed fish these apparent along important with U.S. on and fish from of volunteers research and partners partners Phoenix and stocks. the Phoenix monitoring islands assessments, the 2000-2004, may wildlife. and decline wildlife PRIAs: at west minimally-disturbed of be ship Kingman to to appear expeditions The Islands comprehensive have sh/species tradesintheseislands. Islands, responsible fishing at restore improve side resources, of All in groundings the USFWS coral reef now large but and of to are U.S. and shing. Guam, ecosystems. fishes be two the completed reef surveillance the remaining fish unknown, monitoring to basically will Line for Palmyra assess equatorial of ecosystems the continue conservation densities and the the vigorously refuges, American and 10 NWRs corals substantially species impacts but healthy NWRs Phoenix and but thus As to islands (e.g., two NWRswill believed they be part using of appears in enforcement Samoa, defend far plans inventories , sources of the the with with gray , of are residual Islands, several (Jarvis, several PRIAs PRIAs higher a Areas for to large coral all also new reef and low be all of of page 369 Pacific Remote IslandSidebar Areas The State of Coral Reef Ecosystems of the Pacific Remote Island Areas

OVERALL CONCLUSIONS AND RECOMMENDATIONS Cooperative biennial surveys should continue at these remote islands to: 1) improve the scientific understanding of these fish and wildlife resources as a basis for sound management; 2) monitor spatial and temporal changes in fish and wildlife assemblages in response to natural and anthropogenic forces; 3) help elucidate associations

Areas between fish (the primary resource of interest) and other components of the coral reef ecosystem; and 4) assist the USFWS and other resource managers to improve the scientific basis for protecting fish and wildlife within NWRs and other MPAs. Pacific Remote Island

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