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Short-Range Movements of Hawksbill Turtles (Eretmochelys imbricata)from Nesting to Foraging Areas within the Hawaiian Islands1

Denise M. Parker,2,3,4 George H. Balazs,3 Cheryl S. King,5 Larry Katahira,6 andWilliam Gilmartin5

Abstract: Hawksbill sea turtles, Eretmochelys imbricata, reside around the mainHawaiian Islands but are not common. Flipper-tag recoveries and satellite track-ing of hawksbills worldwide have shown variable distances in post-nesting travel,with migrations between nesting beaches and foraging areas ranging from 35 to2,425 km. Nine hawksbill turtles were tracked within the Hawaiian Islands usingsatellite telemetry. Turtles traveled distances ranging from 90 to 345 km andtook between 5 to 18 days to complete the transit from nesting to foraging areas.Results of this study suggest that movements of Hawaiian hawksbills are rela-

tively short-ranged, and surveys of their foraging areas should be conducted toassess status of the habitat to enhance conservation and management of theseareas.

Ha w ks b i ll s e a tur tle s, Eretmochelys im-bricata (L.), are found in nearshore habitatsin tropical regions of all oceans. Hawksbillturtles are listed as Critically Endangered inthe International Union for Conservation ofNature (IUCN) Redbook (Hilton-Taylor2000) and as Endangered under the Conven-

tion on International Trade in EndangeredSpecies of Wild Fauna and Flora (CITES)(Washington, D.C., 1973) and the Endan-gered Species Act (U.S.A., 1973). The specieshas historically not been commonly harvestedfor consumption in the Pacific because theedible parts can sometimes be toxic (Fussy etal. 2007). Hawksbills in the Caribbean werenot generally harvested for food before the

1900s because of toxicity, but as hawksbillnumbers declined due to harvesting for scutes(Carrillo et al. 1999, Campbell 2003) toxicityalso seemed to decline. McClenachan et al.(2006) suggested that this reduction in toxicitymay be connected to the declines in hawksbillnumbers because of an increased availability

of less-toxic sponges on which hawksbillsprefer to forage. Large numbers of hawksbills were taken in the Pacific, mainly for theirscutes, which are the thickest and most deco-rative of all sea turtle scutes. Items that werefashioned from bonding, shaping, and carvingthese scutes include combs, jewelry, decora-tive art, and tools (Kaeppler 1978, Canin1991, Hainshwang and Leggio 2006). Despiteinternational protection, hawksbills are stillbeing harvested and exploited for curios andfor tortoiseshell, known in Japan as bekko(Canin 1991, MSNBC News Service 2007).Knowledge of hawksbill movements, loca-tions, and habitat usage of critical marine for-aging habitats can aid state and local entitiesto develop effective protection and manage-ment schemes for the species.

Before 1995, few studies had been con-ducted using satellite telemetry to determinepost-nesting movements of hawksbill turtles.Knowledge about population size and distri-bution of hawksbills was obtained mostly

from flipper-tagging and recapture programs.Records of flipper-tag recoveries have indi-

Pacific Science (2009), vol. 63, no. 3:371 382

: 2009 by University of Hawaii PressAll rights reserved

1Manuscript accepted 31 October 2008.2 Corresponding author (e-mail: Denise.Parker@

noaa.gov).3 NOAA, National Marine Fisheries Service, Pacific

Islands Fisheries Science Center, 2570 Dole Street, Ho-nolulu, Hawaii 96822-2396.

4 Joint Institute for Marine and Atmospheric Re-search, 1000 Pope Street, MSB 312, Honolulu, Hawaii96822.

5 Hawaii Wildlife Fund, P.O. Box 637, Paia, Maui,Hawaii 96779.

6 Hawaii Volcanoes National Park, P.O. Box 52,Hawaii National Park, Hawaii 96718 USA.


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cated that many hawksbill populations withinthe Pacific and Atlantic oceans can have long-distance post-nesting movements. In the Pa-

cific, hawksbill turtles on breeding migrationsmoved between Australia and neighboringcountries including Vanuatu, Solomon Is-lands, Papua New Guinea, and Indonesia. Migratory distances between foraging areasand nesting beaches ranged from 368 to2,425 km (Miller et al. 1998, Bell et al. 1999).A flipper-tagged hawksbill that nested in theSolomon Islands was recovered in PapuaNew Guinea after the turtle had traveled anestimated straight-line distance of 1,400 km(Vaughan and Spring 1980). Post-nesting

flipper-tag recoveries from the Caribbeanprovided movement distances of 1,4002,000km between nesting and foraging areas(Groshens and Vaughan 1994, Hillis 1994,Meylan 1999).

Since 1995, there has been increased usageof satellite telemetry worldwide to monitorthe movements of post-nesting hawksbills. Todate, much of the published turtle-trackingresearch has been in the Caribbean (Bylesand Swimmer 1994, Hillis-Starr et al. 2000,

Horrocks et al. 2001, Troeng et al. 2005, vanDam et al. 2008). These telemetry studiesshowed long-distance movements from BuckIsland, St. Croix, U.S. Virgin Islands, to bothNicaragua (1,400 km) and Venezuela (2,000km), as well as short-distance movementsstaying within 300 km of Costa Rica andBuck Island, Virgin Islands. Conversely, sat-ellite tracking of five post-nesting hawksbillsin the Seychelles, Indian Ocean, in 1998showed movements limited to 35175 km(Mortimer and Balazs 2000).

Genetic studies of hawksbill turtles havebeen published mainly on Caribbean popula-tions (Bass 1999, Mortimer et al. 2007). Pa-cific populations are currently being analyzed(Limpus and Miller 2008; NOAA SouthwestFisheries Science Center, unpubl. data).

The Hawaiian Archipelago (U.S.A.) ex-tends for 2,450 km across an isolated regionof the North Pacific (19N, 155W to 28N,178 W ). The hawksbill, known as ea orhonuea in the Hawaiian language, is rare in

the Hawaiian Islands ( Balazs 1978b, Balazs

et al. 1994, National Marine Fisheries Service[NMFS] and U.S. Fish and Wildlife Service[USFWS] 1998). Hawksbill turtles do not

currently nest or reside in the NorthwesternHawaiian Islands. Only one hawksbill hasbeen recorded in that area: a juvenile foundstranded alive at Pearl and Hermes Reef(2005, Pacific Islands Fisheries Science Cen-ter [PIFSC], unpubl. data).

Historically, hawksbills in Hawaii werenot eaten for food because of the toxicityand unpleasant taste of the meat, but thescutes were used for decorative inlay on im-portant objects such as kahili and for medi-cine, bracelets, combs, and fishing hooks

(Buck 1957, Pukui and Elbert 1992, Roseet al. 1993).

Compared with the green turtle (Cheloniamydas), hawksbill sightings and strandingsaround the main Hawaiian Islands are rela-tively uncommon, and the majority of thesightings have consisted of immature turtles(Balazs 1978a, Keuper-Bennett and Bennett2002, Chaloupka et al. 2008). To date, onlytwo flipper-tagged adult hawksbill turtles havebeen recorded away from the nesting beaches.

Both turtles were tagged nesting at PohueBay, island of Hawaii. One was sighted in waters near Kahekili, Maui, a distance of265 km, shortly after the 2005 nesting season(W. Seitz and C. S. K., unpubl. data), and thesecond was sighted shortly after the 2008nesting season near Honoko wai, Maui, adistance of 260 km (W. Seitz, U. Keuper-Bennett and P. A. Bennett, unpubl. data).

The number of adult female hawksbills hasbeen estimated at less than 100, nesting at 20beaches (Katahira et al. 1994, Mangel et al.2000, King et al. 2007; L. K. and W. Seitz,unpubl. data). Previously, Kamehame, a small,remote beach located on the southeasterncoast of the island of Hawaii, was the mainnesting beach in the Hawaiian Islands (Kata-hira et al. 1994). In addition to Kamehame,12 other nesting beaches can be found onthe island of Hawaii. When combined, thesebeaches currently host the majority of hawks-bill nestings in the Hawaiian Islands (Kata-hira et al. 1994; L. K. and W. Seitz, unpubl.

data). Nesting beaches can also be found on

372 PACIFIC SCIENCE . July 2009


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other islands: two on the northeastern shoreof Oahu (Balazs 1978b), one on Molokai(Balazs 1978b, pers. obs.), and four on Maui

(Mangel et al. 2000, King et al. 2007).Hawksbills are known to be spongivores(Meylan 1988), but details about the diet andforaging areas of the hawksbill in Hawaii arestill being studied. However, preliminary re-sults to date indicate that both sponges andalgae are an important part of hawksbill dietsin Hawaii (NOAA PIFSC, unpubl. data).Preliminary results from genetic analyses sug-gest that the Hawaii nesting population in-cludes only a few haplotypes (P. H. Dutton,R. LeRoux, unpubl. data).

In this study, we characterize the move-ment patterns of nine adult female hawksbills within the Hawaiian Islands. Post-nestingmovements to foraging areas are described toillustrate habitat use and offer recommenda-tions for management and conservation ofthis endangered species.

materials and methods

Argos satellite-linked transmitters, ST-3, ST-

14 (1 watt), and ST-20 (model A-1010, 1/2watt), manufactured by Telonics, Inc. (Mesa, Arizona), were safely and securely attached with polyester resin and fiberglass cloth tothe carapaces of nine female hawksbill turtlesfollowing the procedures described by Balazset al. (1996) and Schroeder et al. (2000). Thetwo ST-3 and six ST-14 transmitters eachweighed 750 g and measured 17 by 10 by 3.5cm with the Tygon-protected antenna ex-tending 13 cm from the top of the transmit-ter. Transmitters were programmed witheither a 6 hr on, 6 hr off duty cycle or a 9 hron, 3 hr off duty cycle. Units were turned onat a time computed to synchronize with opti-mum satellite overpass coverage. The singleST-20 transmitter, programmed with a dutycycle of 6 hr on and 48 hr off, weighed 276g and measured 6.0 by 12.3 by 2.8 cm.

After completing nesting and before re-turning to the ocean, each turtle was carefullyconfined in a prone position inside a portableplywood pen while the satellite tag was at-

tached to the carapace. Transmitters were

attached to four hawksbills nesting at Kame-hame on the island of Hawaii (two in August1995 and two in August 1996) and five

hawksbills nesting on the island of Maui atthe following beaches: Kealia (1997, 2005),Kawililipoa (1998, 1999), and Oneloa (2004).Size of the turtles was measured in centi-meters as straight carapace length (SCL)and/or curved carapace length (CCL).

All deployed transmitters sent data topolar-orbiting NOAA satellites carrying Ar-gos receivers (CLS America, http://www.clsamerica.com). Argos locations were calcu-lated by measuring the Doppler shift on thetransmitter signals, which is explained at the

CLS America, Inc. (2007) website http:// www.argos-system.org/. Positional accuracyis defined by Argos as follows (LC locationcode): LC 3, within 150350 m to the givenlocation; LC 2, within 350500 m; LC 1, within 5001,000 m; and LC 0, >1,000 m.LC A and LC B have no assigned value foraccuracy, and LC Z positions were rejecteddue to their calculation with less than twosignals received during the satellite pass. Posi-tions were considered unacceptable if: (1)

they were located on land, (2) the speed trav-eled between two locations was over 5 km/hr,or (3) the position made a turn greater than90 degrees in less than a 24-hr period. Deci-sions for excluding a position were subjectivebased on these criteria.

The best daily location and the great circleequation with the WGS84 ellipsoid were usedto compute distance traveled (Bowditch 1995,Wessel and Smith 1998). Speed (kilometersper hour) was calculated as the distance trav-eled between adjacent positions divided bythe time spent traveling that distance. Speedover the total track was averaged, and themean speed of transit was recorded. Whenavailable, LCs of 0, 1, 2, or 3 were used fordistance and speed calculations; when un-available, estimated distance and speed werecalculated using the beginning and end pointsand the most likely route taken by the turtle, which was indicated by the data for eachtrack. The end of a track was determined ei-ther by the last Argos position or when posi-

tional locations clustered in one general area

Movements of Hawaiian Hawksbill Turtles . Parker et al. 373


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for more than 1 month. The earliest date atan end point was considered the end date fordistance and speed calculations. Home-rangearea in square kilometers was determined forthree of the nine tracks using a simplifiedminimum convex polygon analysis of the for-aging area data scatter, using only LC 2 andLC 3 data. Only three tracks had data suffi-ciently accurate for analysis.

Average dive time in a 12 hr period andtime of last dive were collected and transmit-ted by the ST-3 and ST-14 transmitters. Av-erage dive time and time of last dive wereconverted into minutes using the formulagiven by Argos (data times 2 divided by 60).A mean was calculated for each of these vari-ables to provide an index for each turtle dur-ing the inactive periods (when the turtle was

foraging, resting, or during the internesting

period) and the transit period (defined by thedate the turtle started moving away fromthe nesting beach until the date it arrived atthe foraging/resting area, or the end data).

results

Hawksbill foraging areas in Hawaii based onthese satellite tracking results are shown inFigure 1 along with known hawksbill nestingbeaches. Six of the nine hawksbills moved tothe northeastern shore of the Hamakua Coaston the island of Hawaii. The other three in-dividuals traveled to different island foragingareas. The turtles traveled distances rangingfrom 90 to 345 km, taking between 5 and 18days to complete the transit (Table 1).

The mean duration of the last dive had a

Figure 1. Distribution of foraging areas (indicated by black dots) as determined by satellite tracking of nine hawksbillsfrom 1995 to 2005. Open stars indicate nesting sites. The entire Hawaiian Archipelago is in the inset for scale.



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range of 3477 min during nontransit peri-ods and a range of 535 min during transitperiods. The index range for average divetime during a 12 hr period was 1479 minfor nontransit periods and 519 min duringtransit periods. Home ranges for the threetracks with sufficient data to analyze were es-timated to be 0.5 to 2.0 km2 (G1.0 km2).

Three turtles nesting at Kamehame, islandof Hawaii, two turtles nesting at Kealia,Maui, and one from Kawililipoa, Maui, trav-eled to the Hamakua Coast of the island ofHawaii (Figure 2AF). All turtles followedthe coastlines closely whenever possible. Two of the Kamehame hawksbills traveledin a counterclockwise direction around CapeKumukahi, taking the shortest route aroundthe island of Hawaii to foraging areas onthe Hamakua Coast (Figure 2A,B). One Ka-mehame post-nesting hawksbill traveled in aclockwise direction around the island of Ha- waii, around the southern tip of the island,north along the west coast and then around

the northern tip of the island before arriving

at the foraging area near Paauilo on the Ha-makua Coast (Figure 2C). All three hawksbills that traveled to the

Hamakua Coast from their nesting beacheson Maui took similar paths. They followedthe southwest coast of Maui to KanaloaPoint, at which time they traversed acrossthe Alenuihaha Channel (Figure 2DF). Thehawksbills deflected southward during theirchannel transit. After reaching the coast ofthe island of Hawaii near Kohala or UpoluPoint, the turtles continued in a clockwisefashion around the island to a foraging sitealong the Hamakua Coast.

Three of the turtles tracked did not travelto the Hamakua Coast of the island of Ha- waii. One hawksbill traversed from Kame-hame, island of Hawaii, in a clockwisefashion crossing the Alenuihaha Channel tocontinue along the north coast of Maui to aforaging area on the western side of KahuluiBay, Maui (Figure 3A). Another traveledfrom Kawililipoa, Maui, to a final foraging

area of Pelekunu, Molokai (Figure 3B). The

TABLE 1

Distance and Duration of Post-nesting Hawksbill Movement from Nesting Beaches on the Islands of Hawaii andMaui to their Respective Foraging Areas

Turtle ID Nesting Site

SCL or CCLCarapace

Lengtha (cm)Days

Transmitting

DistanceTraveled

(km)

Numberof Days

in Transit

Mean Speedof Transit

(km/hr)Foraging Site,

Arrival Location

22126-95 Kamehame,Hawaii

88.1 SCL 263 180 10 0.8 Honokaa, HamakuaCoast


83.0 SCL 277 135 8 0.7 Honomu, HamakuaCoast


82.4 SCL 250 275 12 1.0 Paauilo, HamakuaCoast


85.3 SCL 348 345 18 0.8 Kahului Bay, Maui

4802-97 Kealia, Maui 87.5 CCL 142 231 8 1.2 Kuku Pt., Hamakua

Coast4801-98 Kawililipoa,

Maui89.0 CCL 300 172 7 1.0 Waipio, Hamakua

Coast25692-99 Kawililipoa,

Maui89.0 CCL 322 90 5 0.8 Pelekunu, Molokai

19591-04 Oneloa, Maui 88.0 SCL 372 280 14 0.8 Malaekahana, Oahu53751-05 Kealia, Maui 86.6 SCL 312 254 16 0.7 Weloka, Hamakua

Coast

a SCL, straight carapace length; CCL, curved carapace length.



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Figure 2AF. Post-nesting movement of six hawksbill turtles from nesting beaches to foraging areas on the HamakuaCoast of the island of Hawaii. (A) ID 22126: traveled 180 km in a total of 10 days between Kamehame nesting beachto Honokaa; (B) ID 22134: traveled 135 km in 8 days between Kamehame nesting beach to Honomu; (C) ID 24191:traveled 275 km in 12 days between the Kamehame nesting beach and Paauilo; (D) ID 4802: traveled 241 km in 8 daysbetween the Kealia, Maui, nesting beach and Kuku Point; (E) ID 4801: traveled 172 km in 7 days between Kawililipoa

Beach, Maui, and Waipio; (F) ID 53751: traveled 254 km in 16 days between the Kealia, Maui, nesting beach andWeloka.


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Figure 3AC. Post-nesting movement of three hawksbill turtles from nesting beaches to different foraging areas. (A)ID 25695: traveled 345 km in 18 days from Kamehame, island of Hawaii, to Kahului, Maui; ( B) ID 25692: traveled 90km in 5 days from Kawililipoa, Maui, to Pelekunu, Molokai; (C) ID 19591: traveled 280 km in 14 days from Oneloa,

Maui, to Malaekahana, Oahu.


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third turtle used a more circuitous route,completing a 104-km loop returning to nestone last time before heading to her foraging

area near Goat Island, Malaekahana, on theisland of Oahu (Figure 3C).

discussion

Hawaii, as an isolated archipelago in themiddle of the North Pacific Ocean, is situ-ated over 4,800 km from the nearest majorlandmass. Transpacific movements have beendocumented in other sea turtle species such asleatherbacks (Dermochelys coriacea) and log-gerheads (Caretta caretta) via tag returns and

satellite tracking (Resendiz et al. 1998, Polo- vina et al. 2004, Benson et al. 2007). Greenturtles also migrate long distances, but solely within Hawaii. Seasonally, green turtlestravel from foraging areas in the main islandsto mate and nest at French Frigate Shoals inthe Northwestern Hawaiian Islands (Balazsand Ellis 2000). Hawksbills, however, areonly rarely found in the Northwestern Ha- waiian Islands and then only as stranded orinjured, perhaps suggesting pelagic waifs. The

Northwestern Hawaiian Islands are now partof a protected marine sanctuary known as thePapahanaumokuakea Marine National Mon-ument, but very few of the hawksbill nestingsites are protected in a similar fashion. Or-ganizations on Maui and the island of Hawaiihave volunteers monitor nesting beaches; thisis not done on Oahu or Molokai. However,in Hawaii, all sea turtles are protected underthe Endangered Species Act and under Ha-waii State law. Hawaii also has a sea turtlepublic outreach and education system, inwhich both federal and state officials cooper-ate to educate the public on proper viewingof sea turtles.

During the decade of this study, nine post-nesting Hawaiian hawksbills were trackedmoving to four different Hawaiian Islands. Movement distances for these turtles wererelatively short, between 90 and 345 km.These data are comparable with the distancesand time spent traveling by post-nestinghawksbills in the Seychelles (Mortimer and

Balazs 2000). Hawksbills tracked from Barba-dos (Horrocks et al. 2001), Puerto Rico

(Hillis-Star et al. 2000, van Dam et al. 2008), Australia (Miller et al. 1998), and the Solo-mon Islands (Vaughan and Spring 1980)

have shown that the species is capable oflong-distance movements (>2,000 km), yetHawaiian hawksbills were not found to makethis type of movement. The dive durationdata indicate that hawksbills spend moretime underwater during nontransit periods(resting, foraging, and internesting) than when they are actively moving betweennesting and foraging areas. This behavior issimilar to that of green turtles in Hawaii (Ba-lazs 1994, Balazs and Ellis 2000).

On completion of post-nesting movements

to a coastal area in the main Hawaiian Islands,most satellite transmissions continued for 26months, confirming the residency of eachturtle at its foraging area. The end of trans-missions was believed to have usually beencaused by drainage of the transmitter batteryover time or from antenna damage or wearsustained while the turtle was resting in areaswith coral and rocky overhangs. It is difficultto ascertain the exact cause of transmitter ces-sation. Manufacturers provide a theoretical

estimated operational life for batteries usedin the transmitters. However, when a trans-mitter stops giving data within a few weeksto a month of arriving in a foraging area, itis often thought that this is due to antenna wear. Evidence for this assumption comesfrom transmitters that have been recoveredfrom green and hawksbill turtles, where an-tenna wear and missing antennae are appar-ent (Schroeder and Balazs 2000).

Argos positional data based on the Dop-pler shift do not have as precise accuracy asGPS transmitters (minimum of 150 m versus5 m [Yasuda and Arai 2005]). However, thedata are accurate enough for general tracking,and higher-accuracy positions (LC 1, 2, and3) allow a relatively good estimate of the po-sition of the turtle. High-accuracy data were very limited within foraging areas; henceonly three of the tracks could be used forhome range estimation. Based on the datacollected for these three tracks (LC 2 and 3data), estimates of hawksbill home ranges in

Hawaii were calculated to be 0.5 to 2.0 km2

(G1.0 km2). These home ranges were slightly



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larger than hawksbill home ranges found inthe Caribbean area (


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