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Coral Reefs (1995) 14: 237-251
The 1991-1992 rapid ecological assessment of Palau's coral reefs
J. E. Maragosl, C. W. Cook Jr.21 Program on Environment, East-West Center, 1777 East-West Road, Honolulu, HI 96848, USA2 The Nature Conservancy, Pacific Region, 1116 Smith Street, Suite 201, Honolulu, HI 96817, USA
Accepted: 14 August 1995
Abstract. At the request of the Palau and US governments,a team of 30 scientists under the leadership of the NatureConservancy completed a rapid ecological assessment(REA) of nearshore marine resources in Palau in 1992. TheREA provided ecological input to Palau's ongoing masterplan for economic development and identified 45 marinesites worthy of special protection. The REA relied onprevious literature, 1992 aerial photography, interviews,and field observations. A combination of qualitative andquantitative techniques were used to assess stony corals,other reef invertebrates, reef and shore fishes, macroscopicalgae, seagrasses, sea turtles and other marine organisms.The REA covered a variety of coral reef habitats includingb"eaches, seagrass beds, fringing reefs, lagoons, passes,channels, reef holes, patch and pinnacle reefs, barrier reefs,atolls, submerged reefs, mangroves, and "rock" islands.Major stresses to Palau's coral reefs include sedimentationfrom soil erosion, overfishing, and damage from periodicstorms and waves. Minor stresses include dredge-and fillactivities, sewage pollution, anchor damage, tourism use,ship groundings, aquarium fish collecting, and minorcrown-of-thorns lAcanthaster) infestations.
identify and describe ecological areas worthy of specialprotection and conservation. The Nature Conservancyassumed leadership for accomplishing the REA with theassistance of scientists from several institutions from theregion, including several volunteers. Unpublished syn-thesis reports completed in early 1994 present the results ofthe REA for the Southwest Islands (Maragos et al. 1994a)and for the main islands (Maragos et al. 1994b). This papersummarizes the findings of the entire REA. The full detailsof the REA are being published separately.
Materials and methods
The REA involved the follo\\ring steps:.Meetings and interviews with Palau officials and governmentleaders.Acquisition of high-resolution, low-altitude aerial photographs(in April and May 1992).Thorough literature review of all previous scientific investiga-tions in Palau.Interpretation and mapping of ecological areas using the aerialphotographs and maps.Interviews with knowledgeable or interested island residents.including the fishing community.Field work including shoreline and underwater survey~.Analysis and write up of technical reports.Preparation and revision of the synthesis reports -!Introduction
The Palau REA was conceived in 1991 after considerable dialogueand discussion between the representive of The Nature Conservancyin Palau (C. W. Cook) and national government leaders in theBureau of Resources and Development. In addition, state govern-ment officials from Tobi and Sonsorol requested advice on manage-ment of natural resources in the six Southwest Islands. A joint Palaugovernment and Nature Conservancy proposal to conduct an REAfor identifying candidate protected areas and other conservationmeasures was funded by the US Department of the Interior. Thisgrant served as the principal catalyst in gaining the additionalsupport of several other institutions and the volunteer participationof experienced coral reef scientists from the region. Grant funds wereused to cover out-of-pocket expenses for travel, food, board andexpenses associated with the field work, aerial photography, andpreparation of published reports. Key Palauan officials were in-volved in every step and decision in the process outlined above.
In October 1994, the Republic of Palau became an inde-pendent nation in free association with the United States.In anticipation of accelerated economic development, thePalau and US governments and the United Nations De-velopment Program sponsored preparation of a national-level and a series of state-level master economic develop-ment plans. Earlier, the Palau government, with US finan-cial assistance had requested a rapid ecological assessment(REA) of all ecosystems throughout the entire archipelagoin 1992 to provide essential ecological and related environ-mental information for the master planning team and to
Carrespondence to: J. E. Maragos
238
Resource interviews were launched in advance in order to guidethe limited field work to identify conservation problems and oppor-tunities and to identify candidate study sites for evaluation asprotected areas. Interviews were conducted with many Palauansduring visits to Koror (Oreor) and eight outer village and island areas(Des Rochers and Matthews 1992). These intel"Views were crucial in:(1) explaining the purpose of the REA and gaining the necessarycooperation and support for field surveys, (2) jjescribing sites wherefollow-up surveys were warranted, (3) describing past and presentuses and controls over the harvest of coral reef resources, and (4)perceiving problems, conflicts, and opportunities regarding reef con-servation.
Most field sites were selected on the basis of recommendationsmade from interviews, interpretation of aerial photographs, andreview of the literature. Some sites were also selected while in thefield. An effort was made to cover a broad cross section of differentnearshore marine habitats and to investigate all areas identified byothers as important or recommended for protection. Reef areasthought by interviewees to be stressed were also visited. In 1991 and1992, all sandy beach areas in Palau were visited to assess sea turtleand sea bird nesting activity.
Approximately 30 researchers participated in one or more of thefour field expeditions for the REA. The first expedition in June 1991,covered the coral reefs and large embayment of Negeremeduu, off thewest coast of Babeldaob, and several areas. iIl the southern lagoon(Chelbacheb) and eastern lagoon. The second expedition, in June1992, covered the six southwest islands. Thc~ third expedition, inAugust 1992, covered the east coast of Babeldaob, Orak, Ngeream,Malaka1, and Koror Islands and a few southern lagoon sites. Thef(lurth expedition covered northern reefs and atolls, the northernlagoon (Toachel Ngkesol), p°.rtions of the southern lagoon, somemajor islands in the lagoon (Peliliu, Ngerekebesang, Mecherchar,Ulong, Ngeruktabel, etc.), parts of the-west coast of Babeldaob, andAngaur Island just to the south of the barrier reef (see Fig. 1). A totalof 184 field sites were surveyed during the 10 'weeks of field time forthe REA, resulting in fairly complete coverage for the entire archipel-ago of Palau. Important resource surveys coD1pleted since 1990 aresummarized in Table la, b.
Excellent maps of Palau were prepared at a scale of 1:25 000 bythe US Army Map Service during the 1940s and 1950s and by the USGeological Survey during the 1980s. In preparation for the REA,low-altitude color aerial photographs were taken over every islandand coral reef in Palau during April and May 1992. The photographs
were taken mostly at a scale of 1:20 000, with some at larger scales(1:4000-1:8000). The film area for each photograph measured 522cm2, allowing detailed analysi~ <:>f smal~ to large .scale ree~f~atures.Knowledgeable Palauans partICIpated m photo-mterpret4tlon andrecommended many sites worthy of field examination.
Exact details of techniques used during underwater sQrveys va-ried according to the investigator and subject, but all. I involvedunderwater observations recorded during a half hour scui)a dive ateach site. The use of preprinted waterproof forms accelerated therecording of notes underwater. For example, one set of forms in-cluded a list of coral species previously reported from Palau andspace to add new species records. Species present at a site werechecked on the list. At the end of the dive the relative abundance ofeach species was estimated using a five-point scale-dominant, abun-dant, common, occasional, and rare. Space was also available on theforms to sketch a cross-sectional profile of the reef, -describe thecondition of the reef, estimate total percent live coral cover, and makenotes on water visibility, currents, and wave action. On reef slopeseach half hour survey began at a depth of 20 m. Divers then began toascend slowly to the top of the reef. Horizontal distances duringascent averaged 100m in the absence of currents. Otherwise diversdrifted with the current. The half hour time limit was normallysufficient to record all coral species because of the preprinted list ofspecies on the forms. However in some cases a few more lnIi utes was spent on recording species seen during the previous alf hour.
Comparable information was also collected for reef and sh retishes(Donaldson 1992, 1993), other marine invertebrates (Richm nd1991;Birkeland and Richmond 1992; Smith, in preparation), a dmarinealgae (McDermid 1993). Notes were also taken on sea turtle~ encoun-tered during the dives and on oceanographic and water qualityconditions (currents, turbidity, etc.)
One member of the team was responsible for taking underwatervideo at each site. Use of underwater video greatly assisted the entireteam in characterizing each reef site and recording observations.About 5-10min of coverage was taped at each site using Sony Hi8video equipment and tape. The video footage was an importantcheck against earlier reef profile sketches and estimates of relativecoral abundance, especially for dominant and abundant species.Video coverage often revealed a few species of corals not seen by thecoral observer during the surveys.
The maps and aerial photographs were used in calculating thearea and length of selected reef habitats, using planimeters and mapmeasures.
Table 1. a Summary of the technical reports prepared as a part of the 1992 Palau Rapid Ecological Assessment and 1991 Ngeremeduu Bayassessment. b Additional recent technical rep<>rts summarizing the status of other imponant species and areas in Palau
ReferencesSubject
Archaeology (SW Is.)Ciguateraand contaminants (Koror)Freshwater biota (Babeldaob)Seabirds and land animalsMarine invertebrates (other than stony corals)Marine plantsNearshore fishesReefs and stony coralsResource use assessmentsSea turtles
Vegetation (Ngeremeduu Bay and SouthweS1 Islands)
Hunter-Anderson (1993) ::~:.1~ Lobel (1992) i!':.~,
Bright (1991), Smith (1991), Honigman (1992) ~;"~iDi Rosa (1992), Kepler (1993) ,.;
Richmond (1991), Birkeland and Richmond (1992)Gordon (1991), McDermid (1993)Amesbury (1991), Donaldson (1992, 1993)Maragos (1991, 1992a), Maragos and Meier (1993a,lb)Kitalong (1991), Des Rochers and Matthews (1992)Geermans (1992), Di Rosa (1992), Maragos (1992b)jGuilbeaux (1992), Geermans and Honigman (1993)!Canfield et al. (1992), Kepler (1993) ,
Messel and King (1991), Brazaitis (l992)Preston (1990), Kitalong and Oiterong (1991)Marsh et al. (1992)Engbring (1992)~iles and Engbring (1993)Birkeland and Manner (1989t .,.Birkeland et al. (1989) .,.,
-""
Sea water crocodilesNearshore fisheriesDugongsForest birdsBatsNgerukeuid Islands Wildlife PreserveCoastal development sites
239
Physiography
Palau is the westernmost archipelago in the Pacific Oceanand is located at latitudes 6"53' to 8"12' N and longitudes134"08' to 134°44' E (Fig. 1). The main Palau Islands are740 km east of Mindanao and 1300 km southwest ofGuam. The archipelago stretches 800 km from VelascoReef and Ngeruangl Atoll in the north to Helen Atoll andTobi Island in the south. The archipelago consists of 8large, 12 intermediate and 566 small islands (Fig. 1). Themain islands, between Babeldaob and Peliliu Islands, en-compass a distance of about 125 km. Coral reef habitats interms of area, length and type are summarized for each ofthe 16 political subdivisions or states of Palau (Table 2).
The archipelago extends northeast to southwest alongthe Palau ridge, a submerged geological feature stretchingfrom the Mariana and Yap Islands in the north to IrianJayain the south.. All but 11 of Palaul's 586 islands areclustered in the "main" islands. Due to tectonic activity,many of the main islands consist of reels thrust above sealevel, including the rock islands (Chelbacheb), Peliliu Is-land and Angaur Island. Most of the :largest islands arevolcanic, including Babeldaob (332 km 1, Ngerekebesang(5.2km2),Malakal(2.3 km2), and part oJ'Koror(9.13km1.A large deep pass (Toachel Mid) cuts through the mainislands from west to east, and Koror is the only volcanicisland south of the pass. Two low-lying coral atolls(Ngeruangl, Kayangel) and a submerged reef (Velasco)occur to the north of the main islands. Five low-lying coralislands (Sonsorol, Fana, Pulo Anna, Merir, Tobi) one atoll(Helen), and one raised coral island (Angaur) lie between10 and 600 km south of the main islands (Fig. 1).
area (Birkeland and Manner 1990). Surveys of KayangelAtoll were also accomplished in 1974 as part of a ~egionalecosystem survey of the South Pacific (Dahl 19180). Anexcellent inventory of wetlands, including mangro~es, wascompleted for Palau by Stemmermann and Probx (1978).A coastal ecological survey of proposed construction siteson Babeldaob and Koror was also accomplished b~ Birke-land et al. (1989). !
Palau's islands and coral reef ecosystems can Ibe con-veniently divided into seven regions as described~elow.
Northern islands and reefs
Results of observations on corals and reefs
The northern reefs consist of Velasco Reef (a su mergedatoll-like reef), Negeruangl Atoll, and Kayang 1 Atoll.Much of Velasco Reef is uncharted and it exten s 32kmnorth of Ngeruangl Atoll at the southern end of he reef.Velasco is oval shaped and has a maximum est-westwidth of 16 km. The central depression ("lagoon') is 31-55 m deep, while the rim is IS-20m deep. Ngerua gl Atollhas a single unvegetated island, a shallow lagoon with anaverage depth of 6 m, and about 115 pinnacle a d patchreefs. A single tortuous deep pass cuts through the pe-rimeter reef on the southeast side. The lagoon floor iscovered with thick sand deposits and thickets of s aghornAcropora. Heavy wave exposure limits coral dive ityandcover on Velasco Reef.
Kayangel Atoll is located only 3 km north of th north-eastern tip of the barrier reef encircling most of t e mainislands. Beaches off both atolls were important nestingsites for sea turtles but nesting frequency has eclined.Uninhabited Ngeruangl Island is also important or nest-ing sea birds and a few fruit bats previously ro sted onKayangel (Kepler 1993-):-' I
Kayangel has four small vegetated islands, the largest ofwhich (Ngcheayangel) is inhabited. Narrow sea grass bedsand sandy beaches fringe the larger islands. The lagoon hasan average depth of 6 m and a maximum depth of 9.5 m,and about 25 large pinnacles can be detected froPl aerialphotographs. One shallow pass bisects the perimeter reefon the west. Coral diversity and abundance are low in thclagoon. Large fish, dolphins and foraging sea turtles arecommon near the pass.
Northern barrier reef and lagoon
The club-shaped northern barrier reef complex extends31 km north of Babeldaob, enclosing a large lagoon area(Toachel Ngkesol) measuring 325 km2. The lagoondeepens from north (13-21 m) to south (38-50m) and isfilled with many mounds, pinnacles, patch reefs, and reefholes. Two small, high limestone islands (Ngerkeklau,Ngerechur) are situated in the northern lagoon just northof Babeldaob. Expansive seagrass beds are found on thereef flats fringing the lagoon shoreline of northern Babel-daob and extend up to 2 km offshore. Six large deep passes(Kloul Euchel, Telebadel ra Ngkesol, Telebadel raNgerael, Ngamegai, Ebil, and Erengoll-Iengel) cutthrough the northern barrier reef. The passes are progress-
Previous research on coral reefs in Palau is summarized inUNEPjIUCN (1988). For corals, the first observationswere by Rehberg (1892). Later, Japanese marine biologistsconducted reef studies centered at the Palao BiologicalStation near the urban center at Koror. Much of theresearch focused on systematics, physiology, and basicbiology, although some ecological survc~ys were performedat "Iwayama Bay" to the south of the town on Koror,including studies on corals (Eguchi 1936; Abe 1937; Abeet al. 1937; Matsuya 1937; Motoda 193':); 1940a, b). Eguchi(1938) described .170 species of corals from Palau, mostlyfrom Iwayama Bay.
Contemporary research on Palau's corals and reefswere accomplished by visiting scientists from the nearbyUniversity of Guam. One study involv.~d an assessment ofsewage outfall impacts at Malakal (Birkeland et al. 1976),and another focused on a proposed resort atNgerekebesang (Randall et al. 1978). f'aulkner (1974) andFaulkner and Chesher (1979) also published photographicessays of Palau's reef and corals that focused worldwideattention on Palau's spectacular coral reefs and "rock"islands. More ~ecently coral reef surveys have concen-trated in the southern lagoon and rock islands. Reef andterrestrial habitats were surveyed at the Ngerukeuid ("70Islands") Reserve, Micronesia's only marine protected
240
241
Hydrographer Bank
11b
242
0
0
SONSQRCX- ISLANDS
SONSOROL
e
1c
d f 9
Fig. la-g. Maps of the Palau lslands-and reefs, showing shorelines(bold lines). shallow reef margins (thin lines), REA field sites in1991-1992 (open circles), approximate locations of proposed pro-tected areas (numbers), place names (lower case), and political subdi-
visions and regions (upper case). a Northern half of main islands;b southern half of main islands; c-g Southwest Islands: c Sonsoroland Fana, d Pulo Anna, e Merir, fTobi, g Helen. All maps orientedwith true north at the top
-ReefHolesNo.
IslandNo."
Totalarea"km2
93.0[31.3138.8154.5
499.16.7.719.210.1.36.464.620.840.42;68.3
33.719.2
1 39.7
55.015.015.022.5
325.023.812.10
23.030.0
500.0000
103.012.1
1136.5
231141616
2404300
8617
683100
8525
1457
00000000000000
57.97.l
65.0
0000000000
0000
61.713.675.3
402
1.41.41.
1.55007
871.00
50020
418
2170022
286.34.07.22.13.41.31.71.87.91.64.90000
45.0
8.22.77.5
13.823.023.25.88.46.9
22.719.235.52.68.37.02
194.8
79.7 57.4
141.5 62.7
22.050.921
41491134225
5858
,
Aimeliik 27.0Ngetpang 189.9 7.1Ngeremlengui 123Ngardma!1 11.0Ngerechelong 20.8 813Ngaraard 17.3Ngiwal 58.5 0Melekeok 0Ngchesar 4.7Airai 4.0 6.8Koror 100.0 86.4Peliliu 0 0Angaur 0 0Sonsorol 0 0Tobi 0 0Kayangel 0 0Total 264.7 135.3
--
.Excludes Babeldaob. which is divided into 10 states. and which would constitute the 586th island in Palau
11.6
12.713.425.5
2.00
334.4
20.1112.8
000
53.89.6
468.4
243
most major ecosystem types found in Palau. The~ f inging
and barrier reef complex contains numerous r holes
(204) and patch reefs (500), and the lagoon is deep t rough-out (25- 50 m). The fringing reef is long (190 km), a is thebarrier reef (44km). The lagoon slopes generally~pporthigher coral coverage and diversity and a great va~ 'ety of
suspension-feeding hard and soft corals. The re f flats
support abundant sea cucumber populations (Ric mond1991). A total of 19 small islands are found along the westcoast of Babeldaob and are mostly of volcanic origin. Onlythree large passes occur along the entire western coast ofthe main islands. The northernmost (Iengel) is 10tn deepand 570 m wide. The second pass Deimechesengel, a fewkilometers to the south, measures 760 m in width and 44 min depth. A strong feeder current system (Real Edeng) runsthe long distance along the lagoon from Deimechesengalto the next pass to the south, Mlengui. To the south ofMlengui, the next pass is 85 km away at Ngemelis. Thewestern passes are among the most important in Palau forfish migrations and they support high diversity and abun-dance of corals and fish. Coral coverage and diversity aremoderate to high on ocean-facing reef slopes.
ivelydeeper and narrower in a counter-cl!Jckwise directionfromSEtoSW. Portions of the northern 1agoon supportedlarge schools of reef fish (Donaldson 19~13) and moderateto high coral diversity and coverage (Maragos and Meier1993b). Manta rays were reported in the passes. Thehighest coral development on the northern reefs occurredalong the steep west-facing ocean reef slopes (60-70%cover). The reef slope is not as steep along the east coast,where well-developed spurs and groove~; occur along theupper margin.
East coast of the main islands
The eastern coral reefs extend mostly off Babeldaob Islandbetween the northern lagoon and the southern lagoon. Inthe north between Ngerechelong and Ngiwal is a fringingreef; this is the only sector around the main islands lackinga barrier reef and lagoon. South of Ngi'Nal to Melekeok,the reef is a transitional type between fringing and barrierreef. Off Ngchesar to Airai a well-developed barrier reefand narrow lagoon complex is found off the eastern coast.The east-west channel Toachel Mid separates the northernand southern reefs, islands, and lagoons.
The coral reef ecosystems are diverse, including exten-sive mangrove fringes south of Ngaraard and white sandbeaches north of Ngaraard. Seagrasses are also extensiveoff Ngiwal and on Ngerechelong and remain abundantalong the rest of the eastern coast. Spur and grooveformations are well developed off the upper eastwardocean-facing reef slopes and are better developed to thenorth. Dugongs were frequently sighted in the beds (Marshet al. 1992). Ocean reef slopes are steep aDd show high coralcover and diversity. Some pass and lagoon reef sites alsoexhibit high coral abundance and diversity and aredominated by suspension feeders. Some damage fromstorms (especially Typhoon Mike in 1990) was evidentalong the northern end of the reefs where coral diversityand cover are lower. Sediment damage from previousairport construction, dock construction and ongoing roadconstruction has also degraded reefs in Airai Bay to thesouth and off Ngchesar and Melekeok to the east. Reefholes are numerous (87), and 40 small, high limestoneislands are found along the coast, mostly near the southernend. Six channels and five passes (Toachel Ngedbaet,Toachel Suul, Ngemelachel, Orakibad, Toachel Mid) cutthrough the fringing and barrier reefs.
Southern lagoon
South of the main east-west channel of the main islands(Toachel Mid) is the southern lagoon and barrier reefcomplex, covering an area of 500 km2, the largest in Palau.Although fringing reef development is low (22.7 km2 areaand 11.6 km length) the barrier reefs are very extensive,covering 100 km 2 and having a length of 86 km along theocean perimeter. The region also has the greatest numberof patch reefs (683) and the greatest number of islands(491), mostly raised limestone islands. The islands supportrare species of plants and birds (Birkeland and Manner1990; Engbring 1992). Reef holes (100) are common, butmangroves are poorly developed and confined to Korol"Island and the northeast side of Pelilu. Seagrass beds arewell developed off the three large islands of the urbancenter in the north (Koror-Malakal-Ngerekebesang) andoff the southern reefs north of Peliliu Island. The entiresouthwestern side of the barrier reef lacks passes and onlythe dredged German Channel near the southern end of thereef at Ngemelis affords subtidal exchange between thelagoon and ocean. The southeast side of the barrier reef hasseveral gaps and passes between the ocean and :lagoon.Immediately southwest of the urban complex are foundnumerous small to large high limestone ("rock") islands,sublagoons, marine lakes, and reefs. The lakes are uniquein lacking a surface connection to the lagoon or ocean. Thelakes number 67 and are found mostly within the interiorof the larger limestone islands (Peliliu, Syngall-Koror,Ulebsechel. Ngeruktabel. Mecherchar). Some lakes sup-port unique aquatic ecosystems (Hamner and Hal1r11981;Hammer 1991). Beaches are small but numerous andsupport the most important nesting habitat for h~wksbillturtles in Micronesia. The beaches are also popul~r picnicsites for residents and visitors. and Palau's most populardiving sites are located on the southern portiop of thesouthern lagoon and barrier reef. Giant clams and fish
Western coast off Babeldaob
The coral reef ecosystems off western Babeldaob supportthe most extensive fringing reefs, mangrove swamps, fresh-water swamps and estuaries in Palau, and the barrier reef,seagrass beds, and lagoon are also extensive. The north-western lagoon is also the most important feeding groundsfor the dugong, an endangered marine: mammal (Marshet a1. 1992) and the salt water crocodile (Messel and King1991; Brazaitis 1992). Ngeremeduu Bay, including sur-rounding watersheds and offshore reefs, is the most promi-nent feature of the coast and includes representatives of
244
south to north) are low coral islands resting atop tablereefs lacking lagoons, and the dominant habitats cQnsist ofreef flats and slopes exposed to open ocean conditi~ns. Allislands except Fana are inhabited; Helen has only beenrecently occupied on a permanent basis. Helen supportsone of the largest sea turtle feeding areas in Palau a$d untilrecently supported high levels of sea turtle nesting activity,which is now declining. Merir still supports the largestgreen turtle nesting activity in Palau, but present sea turtleharvesting levels are not sustainable (Geermans 1992).Helen supports the largest sea bird nesting populations,although there have been recent declines owing to humandisturbance and harvesting. Sea bird populations are alsoabundant and diverse on uninhabited Fana Island (Kepler1993). Coral diversity was very high at Helen Atoll. Else-where in the southwest islands coral species diversity islower. Reef fishes showed similar patterns, with the highestvalues reported at Helen, where coral reef habitat abun-dance is higher and more diverse than all the rertlainingsouthwest islands combined. Illegal or excessive harvest-ing of giant clams has been reported from Helen Atoll(UNEP/- IUCN 1988), and some evidence of overharvest-ing was apparent during field observations. Fuel sJ)ills andship groundings have also been reported recently in theSouthwest Islands.
were previously more numerous but are now heavilyharvested because they are popular delicacies. Many reefpoints, passes (including Denges and Sar) and dead-endchannels off the southwest perimeter (Ngeumekaoul, Re-botel, Ongingiang) are sites of seasonal fish spawningaggregations. Feeding green and hawksbill turtles arenumerous in the southern lagoon and around Peliliu (nearthe seagrass beds). The salt water crocodile is also commonoff Peliliu's mangroves, and feeding dugongs are commonon the seagrass beds off Sar Passage and Malakal Harbor.Nautiloids are also common off the deep steep slopes of thesoutheast perimeter.
Extensive studies were recently conducted in NgerukeuidIsland (2-6 km1, as part of the long-range planning andmonitoring for Palau's only marine protected area(Thomas et al. 1989; Birkeland and Manner 1990). In 1991and 1992, other sites were also surveyed in the southlagoon as part of the REA. Coral abundance and diversityvaries from moderate to high on lagoon slopes and passesto high in the dead-end channels. During a 45-min surveyin Ngermekaoul dead-end channel in 1991, over 90 speciesof corals were recorded with some individual coral colo-nies measuring 25m across and 10m in height. Ocean reefslopes are steep and support moderate to high speciesdiversity and abundance. Off Malakal Harbor the onlyimportant crown-of-thorns starfish aggregation in Palauwas observed; it is situated in the same location as waspreviously reported (Birkeland ~t al. 1976) and has notspread to other areas (C. Birkefand, personal communi-cation).
Discussion
The status of Palau's nearshore marine ecosystems
Before the 1992 REA, coral reef ecosystems in Palau hadnot been well evaluated by scientists except in a few areas,preventing accurate comparative evaluation throughoutthe whole archipelago. By relying principally on qualita-tive techniques, experienced investigators were ,able tosample adequately all regions and all major ha~itats ina total of less than 3 months field time. Interviel.vs withknowledgeable islanders beforehand helped to focus theefforts of investigators at many key locations. Fpr mostsites the characterization by interviewed islanders wereaccurate, informed, and helped scientists to save time in thefield during surveys. The REA and other recent studiesreveal that coral reefs in Palau are in exc,ellent cqndition,supporting diverse and abundant coral reef, seagra~s, man-grove, and lagoon ecosystems. At present only a f~w coralreef areas have been subjected to anthropogenic ~mpacts,mostly from soil erosion attributed to previous air~eld andongoing road construction and some land cleapng, es-pecially along southern Babeldaob Island. The populationof Palau is small (approximately 15 000) and t~o-thirdslive in the urban center of Koror (Oreor) Island. I1>redgingand filling activities have caused some damage !to man-groves, sea grasses and coral reefs around Koror, Malakal,Ngerekebesang, Peliliu, Angaur and Babeldaob islands.However the extent of damage is not great e~cept atNgetpangand southern Ngeremeduu Bay ofthe\'f1est coastof Babeldaob where road, fishpond, and dock con$tructionhave degraded about 15 ha of mangrove and !seagrasshabitat and unspecified freshwater swamp habit~t. Pollu-tion from the municipal sewage outfall and sanitary land-
Peliliu and Angaur
Peliliu (12.7km2) anchors the southern end of the barrierreef, and Anguar (8.4 km2) is situated 1 0.5 km to the south-west in the open ocean. Both are inhabited, and are raisedlimestone islands. Small boat basins were blasted out of thefringing reef off the west coasts of both islands. Sea turtlenesting has declined to low levels offPeliliu's broad west-ern beaches and is also declining off Angaur. Fringing reefsand steep ocean reef slopes are the dominant reef habitats,except off the northeast lagoon shore: of Peliliu, whichsupports extensive mangroves and seagrass beds. Man-groves are absent off Angaur, but a small seagrass bed isfound along the southwest coast. Coral abundance anddiversity is low to moderate off Peliliu, but fish diversityand abundance is very high. Coral coverage was very highalong the western ocean slopes of Angaur, and diversitywas high along the eastern slopes.
Southwest..1 slands
The six Southwest Islands south of Angaur are 340-600 kmsouth of the main islands. The southernmost is HelenAtoll, a true atoll with one deep pass, a lagoon measuring1. 03 km 2 and containing about 85 patcr, and pinnacle reefs,
and one small vegetated low coral islet (Helen Island)measuring 3 ha in area. Elsewhere, the remaining fiveislands (Tobi, Merir, Pulo Anna, Sonsorol and Fana, from
245
Table 3. Criteria used to identify specific ecological areas Jscandi-dates for protected areas. Lctter codes correspond tothosejlisted inTable 5 I
a Critical habitat for rare species: including endemic, rare,threatened, endangered, or depleted species, such as giartt clams,coconut crabs, salt water crocodiles, ses turtles, dugongs, andmegapodes
b Peak habitat for other important species: especially aggregationsites for edible species of fish that are heavily harvested
c Areas of unique species communities or ecosystems: for example,nautiloids, sea snake aggregation sites, marine lakes, and fresh-water lakes (the last of which are rare in Palau)
d Areas supporting sel-'eral to many important ecos}'stems: siteswhere a variety of ecosystems or habitats are concentrated ina smaller area, such as Ngeremduu Bay where eleven majorecosystem types are found in the watershed and offshore waters
e Important ecological areas supporting several to many rare orimportant species: areas where two or more rare or importantspecies are concentrated, for example, the large seabird and turtlenesting sites at Helen Island
f Other important ecological areas already established by nationalstate or traditional laws or controls: for example, many fish aggre-gation and trochus areas are established as conservation areasand reserves. Also included is the Ngerukeuid ("70 Islands")Ecological Reserve, the only established protected area in Palauaffording full protection to all species
g Important ecological areas threatened with development, habitatdestruction, or overharvesting of key resources: for example, man-grove swamps, seagrass beds or coral reefs in the path of destruc-tion from road building, soil erosion, resort development,dredging and filling, and chronic solid waste pollution andsewage discharges
Table 5, Fig. 1). Exact boundaries for these areas were notdelineated since full discussion with the community andresource users had not been completed at the time ofwriting. The REA to datellas focused primarily on marineand coastal ecosystems, although some recommendationshave also been directed toward the protection of uplandspecies and ecosystems. For example, important habitatsfor rare birds, roosting fruit bats, and endemic freshwaterspecies have been proposed for protection. Aside fromrecommending the designation and management of a sys-tem of protected areas, the REA also led to recommenda-tions to control fish harvesting activities and targetedseveral groups of rare species for special attention, includ-ing sea turtles, crocodiles, dugongs, fruit bats, c9conutcrabs, giant clams, top sh~ll, a~d certain bird--ipecies.Many of the recommendations In the REA were basedupon feedback from local governments and islanders dur-ing the interview phase of the assessment. For example, thestates of Kayangel and Tobi advocated greater levels ofprotection for the natural resources at Palau'slhree atolls:Helen, Ngeruangl, and Kayangel. State governments alsorequested greater cooperation with the national ~overn-ment in enforcing local conservation ordinances land inrespecting the customary tenure and resource ~anage-ment regimes which have operated successfully o"er mil-lenia to sustain the productivity of harvested spec~es andhabitats. However on some remote islands (T~bi forexample) much of the population has moved to the; Kororurban center and many traditional practices and! tenureare no longer followed.
fill disposal areas in the Koror urban area may also bedegrading water quality and nearshore habitats, althoughit is difficult to document the extent of l:he impacts. Shipgroundings and oil spills are always a threat, especially atthe low coral island and reef areas; a 1992 ship groundingand spill at Helen Atoll appeared to res1ult in only minordamage.
Although marine resources in Palau appear to be inrelatively good condition at the regiona!ljecosystem level,fishing and harve~ting activities have reduced many spe-cies, and populations of several species, including dugongs,nesting sea turtles, and crocodiles, are in danger of beingextirpated from Palau. In the 1970scrocodiles were huntedfor their skins, and dugongs have btoo-en traditionallyhunted for their meat. The populations of both specieshave dwindled to very low levels. The green turtle is huntedfor its meat and eggs. The rare hawkbill turtle is hunted forits eggs and shell. The latter is fashioned into jewelry that issold in stores on Palau. Other groups are also heavilyharvested including depleted "food" fish, (particularly thebump- headed parrotfish and Napoleon wrasse), trochus,giant clams, coconut crabs, and mangrove crabs. Aquar-ium fish and "food" fish are also harvested in heavynumbers and shipped to nearby overseas markets. Al-though fruit bats are no longer exported, they were heavilyexported from Palau to the Mariana Islands during thepast decade.
The greatest threat to nearshore marine ecosystems inPalau stems from the changing political status and econ-omy of Palau. The 1994 transition from a trust territory toan independent nation means that US environmental lawsand regulations no longer apply to Palau. Furthermore,the new political-economic relationship between the USand Palau, termed the Compact of Free Association, willprovide additional sources of funds for capital improve-ment and economic development. The relaxation of USjurisdiction is also beginning to fuel large-scale economicdevelopment in Palau from foreign investors, especially onBabeldaob, which is relatively sparsely inhabited. Huntingand harvesting of threatened and endangered sea turtles isnow increasing. Construction to expand airport capabili-ties, complete the new capital and expand the road net-work on Babeldaob are also leading to land clearing andpotential soil erosion, which is causing sedimentation incoastal waters. Fortunately, most of Babeldaob is en-circled by an intact mangrove fringe which serves asa protective barrier, impeding sedimentation in coastalwaters. However several swamps have t>een degraded ordestroyed by clearing, excessive soil deposition, and land-fill expansion for docks and urban development. Clearly,planning for future development and conservation isneeded to maintain the high quality and diversity ofPalau's coral reef and related ecosystems.
The preliminary results of the REA (Maragos et aI.1993, 1994) have been passed on to Palau governmentleaders and the preparers of Palau's new master plan foreconomic development. The REA led to development ofcriteria and categories of protected area!; (Tables 3,4) andto the identification of about 45 marine ecosystem areaswarranting specialprotection due to a combination of highvalue and/or imminent threat to habitats or species (see
246
Table 4. Types of ecological protected areasrecommended for designation in the Palau1992 REA
National park
Ecological reserve
Forest preserve
Marine preserve
Coastal conservationarea
Fishery conservationarea
Tourism site
Special managementarea
Containing a combination of highly significant cultural, historical,esthetic, natural, or ecological attributes and resources of nationalor international significance. Access controlled through,permitsand no harvesting or collecting of any type allowed. Activelymanaged (staff on site)
Area containing unique or important combinations of terrestrial,freshwater, estuarine or marine habitats and species
Area supporting unique or important forest or upland habitat andspecies of plant and wildlife
Area supporting unique or important marine habitat or species ofplants and animals, especially breeding or aggregation site forrare, unique or important species
Shoreline or beach area supporting important ecologicalhabitat for vegetation and wildlife
Limited entry area for fishery of recreational,subsistence or commercial significance warranting special protection or controls to promote sustainable use and avoid over-harvesting and habitat destruction
Visitor destination site with pristine and esthetic value potentiallypopular for visitation and recreation where special controls areneeded to maintain sustainable multiple uses
Area proposed for large-scale development (transportation, resort,residential subdivision, golf course, utility) where importantnearby ecosystems or species may be threatened unless thedevelopment is properly planned and managed
logical scientists to contribute to the conservation of im-portant ecosystems and species, especially where time,funding and geography pose constraints on more orderlyscientific scrutiny and research. For example, there arevery few coral reef scientists in the many island groups inthe tropical Pacific, and the great majority of reef areashave never been visited by scientists (Maragos et al., inpreparation). Fewer scientists are actively engaged in coralreef management activities such as monitoring, pollutionimpact assessment, restoration, protected areas manage-ment, coastal zone management, and rapid ecologicalassessment. Most of these scientists are focusing on coralreefs near urban areas where logistical problems are moreeasily solved, where a variety of stresses to reefs are concen-trated (e.g. solid waste and sewage pollution, industrialdischarges, coastal construction, overharvesting of fish-eries), and where political will and funding for research isconcentrated. As a result, scientific resources are verylimited and most often do not anticipate and cannotrespond to the need for scientific scrutiny and input to helpguide future economic development in a manner thatavoids or reduces costly impacts on important ecosystems.
In this study the participation of outside scientists hadto be limited and efficiently organized as a result of timeand funding constraints. Nevertheless, considerable leadtime was needed by The Nature Conservancy in Palau tobuild up rapport and respect to convince island leaders ofthe potential value of an REA. This was especially import-ant because all management and protection of Palau's eco-systems will be borne by Palauan residents and leaders. Inthe case of Palau, the REA was designed to cover the entirearchipelago and provide recommendations to gove~nmentofficials and consultants working on the master plan foreconomic development, and for village-based resourceusers who share marine resource management responsi-bilities with the government in Palau. The short study
The final draft of Palau's master plan for economicdevelopment has been completed and circulated for reviewto decision-makers and the general public. Roughly 90%of the REA recommendations were incorporated into thisfinal draft, and upon the document's approval, imple-mentation of this section should begin sometime in 1995.For the purposes of an independent review of the REAfindings, the master plan's environment1u consultant sur-veyed several state officials in Palau to ascertain whatnatural areas they believed to be important for futureprotection efforts. There was some 75% overlap in priorityarea identification between the states' suggestions andthose of the REA. Provided that a system of protectedareas is formally designated in Palau, their monitoring andon-site enforcement will be crucial for maintaining impor-tant ecosystems and many heavily harvested species.
The REA to date also led to other recommendations toimprove the environmental impact assessment (EIA) pro-cess for development planning, to initiate a long-term coralreef monitoring program, and to provide special technicaland educational assistance to better design, site, and con-struct special types of development, particularly roads andresorts. Most importantly, the preliminary results of theREA were completed in time to be available at the begin-ning of the process to develop the master plan for econ-omic development. The authorization and designation ofprotected areas in that plan and proper execution of EIA,conservation, and enforcement programs should contrib-ute measurably to the sustainable use of Palau's nearshoremarine resources.
The role of RE.4 in coral reefmanagement
Rapid Ecological Assessment represents an int:reasinglyimportant technique and opportunity for social and bio-
-Fishery conservation areaEcological reserve
Velasco Reef.(all) 1Ngeruangl Atoll (all) 2 a,b,g
a,e
Kayangel Ato.ll (Ulach) 3 b.c.e,g
All marine habitats and species to 100m depthNesting wildlife, all marine species, island and
coral reef ecosystems IAll marine habitats and species in channrlFishery conservation area.
tourism siteCoastal conservation area e,g Nesting sea birds, megapodes, sea turtles, fruit
bats, coconut crabsAll harvested speciesFishery conservation area b,c,f,g
Coastal conservation area a Nesting sea turtles
Marine preserve Feeding dugongs and sea turtles
EC()logical reserve c Sea snake aggregations
ca, b,c, d, e, f, g
Marine preserveEcological reserve,
special management areaEcological reserve a,b
Marine preserveEc<)logical reserve,
fishery conservation areaFisllery conservation area,
marine preserveForest preserveMarine preserve,
special management areaF OJ:est preserveForest preserveCoastal conservation area
b,cb,d,f,g
a,e,f
Feeding dugongs and sea turtlesLake biota, crocodiles, rare waterbirds, fruit
batsFruit bats, nesting sea turtle, marine lake
Mangrove and fisheries habitatWatersheds, estuaries, offshore marine habitats
and speciesDugong and sea turtle feeding areas
a,ba,b
Forest and endangered birdsDugong and sea turtle feeding areas
a,b,c,da,b,c,d,ea,c,g
Marine lake, forest, bat and bird habitatMarine lake, forest, bat and bird habitatSea turtle, megapode nesting, forest habitats
National park
Marine preserve
a, b, C, d, e, fga,e
Marine preserve a, b,c,d.e
Kayangel Atoll (Ngeriungs andNgerebelas) 4
Northern lagoon (all) 5(Ngerechelong)
Ngerchur and Ngerkeklau Islandbeaches (Ngerechelong) 6
Ngerechelong and Ngaraad seagrassbeds 8
Yasumba "snake house" andsurrounding reef (Ngaraard) 9
Ngiwalseagrass beds 12Melekeok-Ngiwal forests, lakes, and
rivers (Ngerdok) 14Ngerduais and Ngeream Island
clusters (Airai) 19Aimeliik 43Ngeremeduu Bay 45
Ngardmau and Ngeremlenguimangroves 47
Ngardmau savanna 48Ngemelachel harbor, Ngederrak
Reef seagrasses (Oreor) 34Ngeruktabel Island (Oreor) 35Mecherchar Island (Oreor) 35Ulong, Omekang, Kmekumer, and
NgkesiiUsland beaches 36Rock Islands Conservation Area
(Oreor) 37gar and Denges passes, seagrass
fiats 23,24 .
Peliliu mangrove and seagrass area38
Peliliu forests 39 Forest preserve ornational park
Coastal conservation area
b,e,f
Angaur beaches 41 a,b
Barrier reef passes and (many)channels 7, 10, 11, 13, 15, 16, 1718,20,21,22,23,25,26,27,46,47
Barrier reef points-28,29, 3O, 31,32,33,40,44
b,e,f,gFishery conservation area
Sea turtles, megapodes, tourist sites, giant clams,coral reefs; marine lakcs, forests, etc.
Dugong and sea turtle feeding grounds
Seagrass, mangroves, crocodiles, waterbitd~sea turtles
World War II battleground, coconut crabs, birdpopulations
Nesting beaches for sea turtles, megapodes,coconut crabs
Spawning grounds, aggregation sites for fish ofcommercial and subsistence importan~, highcoral areas, giant clams
Spawning grounds, aggregation sites for fish ofcommercial and subsistence importan~, high.coral areas, giant clams
Coral and fish-rich areas
Fishery conservation area b,e,f,g
Marine preserve,coastal conservation area
Fishery conservation area
f,gNgemelis-Peliliu sport diving andfishing sites 40
Lkes shoal (Angaur-Peliliu) 42 a,b,g
and with the goal of establishing an ecological basel~ne forlongterm monitoring, the field procedures woul4 haveincorporated quantitative transect-type surveys at perma-nently marked sites. In fact, a monitoring program is beinginitiated in Palau for coral reefs, based upon the feedbackfrom the REA.
time, limited funding, and large geographic coverage of theREA only allowed for qualitative and semi-quantitativefield procedures, but the approach was appropriate for themain objective of the REA to focus attention on identifica-tion of a set of candidate protected areas throughoutPalau. Under more lenient funding and time constraints,
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Nevertheless, quality control is a concern for REA: thatrelies heavily on qualitative techniques and observat~ons.Under such circumstances it is essential that experehcedsenior scientists actively participate and direct field! sur-veys and interpretation of results. Comparing the results ofqualitative surveys with the results of "spot" quantitativesurveys could also serve as a check on quality contro~ andquality assurance.
Ideally, a combination of qualitative and quantitativetechniques should be planned for rapid ecological assess-ments, provided that the study objectives and funding andtime requirements are met. Quantitative surveys at fixedlocations are required to assess long-range trends at speci-fic sites, but a more extensive qualitative survey before-hand would help in selecting the specific sites for long-termmonitoring.
The concept of REA of coral reef environments is notnew and many such assessments have been accomplishedto date, although addressing different study objectives(Table 6). For example, the manta tow survey techniquehas been widely used in coral reef environments, especiallyto assess and map crown-of-thorns starfish densities andlive coral cover, and for these purposes the technique iswell suited. In fact UNEPjAIMS (1993) has advocated themanta tow technique as part of a standardized methodol-ogy to monitor coral reefs worldwise. The advantage ofthis technique is broader area coverage per unit of obser-vation time plus the collection of data in a format useful formapping. On the other hand, the manta tow technique byitself is not particularly useful for assessing species diver-sity on coral reefs and for more precise estimates of coraland fish abundance. In order to increase the accuracy ofabundance and diversity estimates, more time must bespent per unit reef area;..-Given that the number of ob-servers, time, and money will always be constraints incollecting information during underwater surveys, it isessential that the study objectives be clearly defined inadvance to determine the appropriate mix of personneland procedures. For this reason, the development ofa standardized set of techniques for surveying coral reefsmust take into consideration that the study objectivesshould dictate the methodology, and that there is noone technique that will serve all purposes. Standardizationof methodology should concentrate on formulating asuite of techniques to address a range of different studyobjectives, and scientists should not be preoccupied withselecting a single technique to address all objectives (seeTable 6).
The techniques for long-term monitoring of coral reefs,for examble, would probably include quantitative tran-sect type surveys at fixed and preferably permanentlymarked sites. However, selection of the monitoring sites isof critical importance, and a prior survey involving othertechniques (qualitative observations) would be ne4ded,especially for reefs where there is little in!ormation o~ thedistribution and abundance of the organIsms and ha~ltatsto be monitored. REA could be designed as the initialphase of selecting and establishing a network of monitor-ing sites. The initial REA could cover a greater area andlead to decisions on which sites should be (or should notbe) included in a long-term monitoring study. If the deci-
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250
sion were made quickJy, the monitoring phase of thesurvey couJd then be accompJished immediateJy after com-pletion of the initial assessment, even during the same fieldexpedition.
the late David Idip, and Stephen Patris on behalf of the Gov~rnmentof Palau; Chuck Birkeland, Bob, Richmond, Steven Amesbury, andBarry Smith of the University of Guam; Karla McDermid, LaniStemmermann, and Kim Des Rochers of the University of Hawaii;Phil Lobel of Woods Hole Oceanographic Institution; LucianaHonigman of The Nature Conservancy; Robert Myers of t~e Gov-ernment of Guam; Robert Smith, Joan Canfield, and Derral Herbstof the US Fish and Wildlife Service; Suzie Geermans of the SouthPacific Regional Environment Program; and Jack Noble and VickiGetz of Greenpeace, Pacific Campaign. The Palau rapid ecologicalassessment was administered and facilitated by The Nature Conser-vancy's Micronesia and Hawaii Field offices and primarily financedby grants from The Office of Territorial and Internaticnal Affairs ofthe US Department of the Interior, the John D. and Catherine T.MacArthur Foundation. KEIDANREN, Greenpeace Pacific Cam-paign, The Nature Conservancy, The South Pacific Regional Envi-ronment Program, and the East-West Center. The authorsacknowledge the institutional support arranged by Jeanne Kirby andPia Mancia of Greenpeace and the strong leadership of NoahIdechong, David Idip. Stephen Patris, and Demei Otobed of theRepublic of Palau. Mary Ha yano of the East-West Center, Programon Environment assisted in the preparation of the manuscript, andKarin Z. Meier of CORIAL assisted in the preparation of the tablesand maps.
Conclusions
References
Rapid ecological assessment is an important technique forcoral reef management and can help guide future manage-ment options, especially on reefs where there is very littlepublished or unpublished information. REA can serve asthe initial step and for setting priorities in developingmarine protected areas, coral reef monitoring, assessinginfestations of Acanthaster predation, damage assessmentfollowing natural disasters (e.g, typhoon~;, waves, sea levelchanges), following anthropogenic catastrophes (e.g, shipgroundings, oil spills), etc. Aside from an initial appraisal,REA is also well suited in situations where the large size ofa reef study area, complex logistics, limited time, andlimited funding support severely restrict the total effortthat can be expended. An essential rOll: for experiencedcoral reef scientists involved in coral reef management andconservation is to participate in REA and train others onthe basic techniques to accomplish REA.
No standardized or detailed methodology is advocatedhere for REA, although the following sequence or stepsshould be considered: -(a) Define with resource managers and users the purposeand objectives of the REA(b) Define geographic scope, based upon the objectivesand constraints(c) Select REA team and assign responsibilities(d) Check with government leaders and other potentialusers-will the REA meet their needs'?(e) Conduct review of literature, maps, and aerial photo-graphs(f) Conduct interviews with knowledgeable residentresource users (fishermen, village elders, historians,curators, other scientists, etc.)(g) Select field sites, relying heavily on the above inputsand steps(h) Schedule and accomplish field worll: and include con-tingency plans for unscheduled disruptions(i) Each participant analyzes results and prepares preli-minary technical report for review by REA leadershipG) Leader(s) prepares preliminary synthesis report andsend to other team members and user i~roups for reviewand comment(k) Finalize and submit synthesis report including re-commendations(1) Finalize and publish technical reports(m) Follow-up with users on subsequent study phases(monitoring, protected area planning, etc.)
Acknowledgements. The REA would not have been possible \\'ithoutmany scientists volunJeering their time to participate in field workand prepare reports: Terry Donaldson, Kay Kepler, RosalindHunter Anderson. Karin Z. Meier. Elizabeth Matthews. MichaelGuilbeaux. Roger DiRosa. Greg Bright. Greg Gordon, and HeidiHirsh (all without compensation): Noah Idechong, Demei Otobed.
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Amesbury SS (1991) Marine, fishes in the Ngerrneduu Bay area,Palau. University of Guam, Marine Laboratory. Prepared forThe Nature Conservancy, Pacific Region, Honolulu
Bass DK, Davidson J., Johnson DB, Miller-Smith BA, Munday CN(1989) Broadscale survey of crown-of-thorns starfish on the GreatBarrier Reef, 1987-1988. Aust Inst Mar Sci, Townsville
Birkeland C, Manner H (eds) (1989) Resource survey of the Ngeru-kewid Islands Wildlife Preserve, Republic of Palau. South PacificRegional Environment Programme, Noumea
Birkeland C, Richmond RH (1992) Rapid ecologicalasses$ment ofPalau-invertebrates. University of Guam. Marine Laboratory.Prepared for The Nature Conservancy, Pacific Region, Honolulu
Birkeland C, Tsuda RT, Randall RH, Amesbury SS, Cushing F(1976) Limited current and underwater biological surveys of aproposed sewer outfall site on Malakal Island, Palau. Universityof Guam, Marine Lab Tech Rep 25:1-59
Birkeland C, Richmond RH, Matson EA, Holthus P, Con~ard NS(1989) Palau coastal resources survey (phase 1: Babeltnuap andKoror-selected sites). South Pacific Regional Environ~ent Pro-gramme, Noumea
Brazaitis P (1992) Recovery plan for the saltwater dToc~ile, Cro.cody/us porosus in the United States Trust Territory of~e PacificIslands, Republic of Palau (draft). US Fish and Wildlife Service,Portland
Bright G(1991) The freshwater environment in the Ngerem~duu Baywatershed. Republic of Palau. Prepared for The Nature Con-servancy, Pacific Region, Honolulu
Canfield JE, Herbst DR, Stemmerrnann RL (1992) Rapid efologicalassessment of areas in the Republic of Palau under cons~derationfor conservation: Ngeremeduu Bay drainage area v~getationreport. US Fish and Wildlife Ser\1ce, Honolulu. Prepare~ for TheNature Conservancy, Pacific Region, Honolulu
Carpenter RA, Maragos JE (1989) How to assess envir<[lnmentalimpacts on tropical island and coastal areas. EnvirontPent andPolicy Institute, East W.est Center, Honolulu
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