forage and migration habitat of loggerhead (caretta

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Forage and migration habitat of loggerhead (Caretta caretta) and olive ridley (Lepidochelys olivacea) sea turtles in the central North Pacific Ocean Jeffrey J. Polovina 1 , George H. Balazs 1 , Evan A. Howell 1 , Denise M. Parker 2 , Michael P. Seki 1 and Peter H. Dutton 3 1 Pacific Islands Fisheries Science Center, National Marine Fisheries Service, 2570 Dole Street, Honolulu, HI 96822-2396 2 Joint Institute for Marine and Atmospheric Research, University of Hawaii, 8604 La Jolla Drive, La Jolla, CA 92037 3 Southwest Fisheries Science Center, National Marine Fisheries Service, 8604 La Jolla Drive, La Jolla, CA 92037 N A T I O N A L O C E A N I C A N D A T M O S P H E R I C A D M I N I S T R A T I O N U . S . D E P A R T M E N T O F C O M M E R C E ABSTRACT Satellite telemetry from 26 loggerhead (Caretta caretta) and 10 olive ridley (Lepidochelys olivacea) sea turtles captured and released from pelagic longline fishing gear provided information on the turtles’ position and movement in the central North Pacific. These data together with environmental data from satellite remote sensing are used to describe the oceanic habitat used by these turtles. The results indicate that loggerheads travel westward, move seasonally north and south primarily through the region 28–40 N, and occupy sea surface temperatures (SST) of 15–25 C. Their dive depth distribution indicated that they spend 40% of their time at the surface and 90% of their time at depths <40 m. Loggerheads are found in association with fronts, eddies, and geostrophic currents. Specifically, the Transition Zone Chlorophyll Front (TZCF) and the southern edge of the Kuroshio Extension Current (KEC) appear to be important forage and migration habitats for loggerheads. In contrast, olive ridleys were found primarily south of loggerhead habitat in the region 8–31 N latitude, occupying warmer water with SSTs of 23–28 C. They have a deeper dive pattern than loggerheads, spending only 20% of their time at the surface and 60% shallower than 40 m. However, the three olive ridleys identified from genetics to be of western Pacific origin spent some time associated with major ocean currents, specifically the southern edge of the KEC, the North Equatorial Current (NEC), and the Equatorial Counter Current (ECC). These habitats were not used by any olive ridleys of eastern Pacific origin suggesting that olive ridleys from different populations may occupy different oceanic habitats. 150E 160E 170E 180 170W 160W 150W 140W 130W 120W 5N 10N 15N 20N 25N 30N 35N 40N 45N Loggerhead Olive Ridley Figure 1. Track lines of 26 loggerhead (dashed) and 10 olive ridley (solid) turtles based on daily Argos reported positions from the period 1997–2001. Solid dots mark the start of each track line. Telonics ST-10/ST-18 position transmitters were attached to 32 juvenile olive ridley and juvenile loggerhead Turtles from Hawaii-based longline fishing vessels from 1997-2001, with an additional 2 olive ridley and 2 loggerheads turtles instrumented with SDR-T10 dive recorders. The spatial distribution of all track lines for the 36 turtles released with transmitters spanned an area from latitude 6–41 N and longitude 130 W–150 E (Fig. 1). Loggerheads were predominately north of about 28 N occupying the northern portion of the Subtropical Gyre and the Transition Zone, while the olive ridleys were generally south of 28 N in the Subtropical Gyre and the equatorial region (Fig. 1). All the loggerheads traveled westward with the exception of two on the eastern edge of the area that traveled southeast, while olive ridleys exhibited both eastward and westward movements. Figure 2. Time-at-depth, dive depth, and daily maximum dive depth cumulative frequency distributions for all time periods combined: (a) both loggerheads combined, and (b) both olive ridleys combined. From satellite SST estimated for each turtle position we estimated that olive ridleys primarily occupied a 5 C SST range, 23–28 C during all quarters, but within that range they appeared to be found centered at 24 and 27 C. Loggerheads occupied cooler water and covered a broader 10 C SST range, 15–25 C, occupying cooler temperatures in winter and spring and warmer temperature in the summer. Both loggerheads exhibited relatively shallow dives with cumulative time-at-depth frequency distributions indicating that they spent 40% of their time at the surface and about 90% of their time at depths <40 m (Fig. 2). Both olive ridleys exhibited deeper diving than the loggerheads, with pooled cumulative time-at-depth frequency distributions indicating that they spent about 20% of their time at the surface and about 40% of their time deeper than 40 m (Fig. 2). 160 140 120 100 80 60 40 20 0 Depth (m) 0.0 0.2 0.4 0.6 0.8 1.0 Cumulative Frequency Time At Depth Dive Depth Daily Maximum Depth B. Olive Ridley 160 140 120 100 80 60 40 20 0 Depth (m) 0.0 0.2 0.4 0.6 0.8 1.0 Cumulative Frequency A. Loggerhead Time At Depth Dive Depth Daily Maximum Depth 0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 0.4 0.6 0.8 1.0 Temperature-depth profiles indicated important differences between the thermal structure in the habitat used by the two turtle species. The olive ridley habitat was characterized by a deep thermocline at 100 m and minimal horizontal surface temperature fronts (Fig. 3). In contrast, the loggerhead habitat was characterized by a shallow thermocline at 50 m and several strong horizontal surface temperature fronts at 20 and 17 C located at 28 and 32 N, respectively (Fig. 3). Similar depth differences in the chlorophyll-depth profiles were also observed, with the subsurface chlorophyll maximum above 100 m in the loggerhead habitat and reaching the surface at 33 N when it became the TZCF, a key feature for loggerhead habitat (Fig. 3). In the region of olive ridley habitat between 12 and 22 N latitude, the chlorophyll maximum was deeper than 100 m, although it shoaled to the surface at 10 N latitude at the northern edge of the NEC (Fig. 3). Figure 3. Temperature depth and chlorophyll depth plots for (a) olive ridley habitat (10–22 N latitude, April 21–30, 2001), and (b) loggerhead habitat (25–34 N latitude, April 24–May 1, 1998). 0.2 0.2 0.2 0.2 0.2 0 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.6 0.1 0.1 0.1 0. 0.1 0.2 2 0.2 0.2 0.2 0.2 0.2 0.2 0.3 11 11 12 12 12 13 13 13 13 13 14 14 15 15 16 17 17 18 18 18 18 19 19 19 20 21 22 10 10 11 12 13 13 14 14 15 15 16 17 18 19 20 21 22 24 24 25 25 25 25 26 27 23 20 40 60 80 100 120 140 160 180 200 26 28 30 32 0.2 0.4 0.6 0.8 Latitude ( o N) Depth (m) 20 40 60 80 100 120 140 160 180 200 Depth (m) 10 12 14 16 18 20 22 Latitude ( o N) 0.25 0.2 0.15 0.1 0.05 0 26 28 30 32 Latitude ( o N) 10 12 14 16 18 20 22 Latitude ( o N) 50 100 150 200 250 300 Depth (m) Depth (m) 30 25 20 15 10 50 100 150 200 250 300 24 22 20 18 16 14 12 10 A. Olive Ridley Habitat - Temperature o C B. Loggerhead Habitat - Temperature o C C. Olive Ridley Habitat - Chlorophyll a (mg m -3 ) D. Loggerhead Habitat - Chlorophyll a (mg m -3 ) Chlorophyll a (mg m -3 ) Chlorophyll a (mg m -3 ) Temperature ( o C) Temperature ( o C) One interesting loggerhead track line shows the turtle’s westward trajectory just south of the strong eastward KEC, which is associated with the sharp gradient in SSH (Fig. 4). The turtle’s westward movement was often assisted by the westward flow on the north side of the counter-clockwise rotating eddies it encountered along 33 N (Fig. 4). During February–April 2001, the loggerhead position, as indicated by the black dots on the track line, was along the edge of a KEC meander. During February, March and April this meander developed into a cyclonic eddy, and the turtle occupied the edge of the eddy (Fig. 4). These meanders and eddies represent regions of enhanced surface chlorophyll as seen from surface chlorophyll estimated from SeaWiFS data for February 2001 (Fig. 5). The two cyclonic eddies at 165 and 180 E and the meander at 170 E seen in the altimetry data for February 12–22 are regions of enhanced surface chlorophyll (Fig. 5). The loggerhead remained for 3 months along the edge of the meander and eddy at 170 E which represents surface convergence zones and a likely site of concentrated prey for the loggerhead (Figs 4 and 5). 160 E 165 E 170 E 175 E 180 175 W 170 W 27 N 30 N 33 N 36 N 39 N 130 140 150 150 150 150 160 160 170 170 170 170 180 180 180 180 180 190 190 190 190 190 190 190 200 210 C. April 12 - 22 2001 30 cm/sec 160 E 165 E 170 E 175 E 180 175 W 170 W 27 N 30 N 33 N 36 N 39 N 140 150 150 160 170 180 180 180 180 190 190 190 190 200 200 B. March 13 - 23 2001 30 cm/sec 160 E 165 E 170 E 175 E 180 175 W 170 W 27 N 30 N 33 N 36 N 39 N 110 140 150 160 170 170 170 180 180 180 180 190 190 190 190 190 200 200 200 210 A. February 12 - 22 2001 30 cm/sec 90 150 160 170 180 190 250 Sea level (cm) B. SeaWiFS Ocean Color, February, 2001 0.000 0.100 0.200 0.300 0.325 0.350 0.375 0.400 0.500 2.000 27 N 30 N 33 N 36 N 39 N A. Sea Surface Height and Geostrophic Currents, Feb 12 - 22, 2001 160 E 165 E 170 E 175 E 180 175 W 170 W 27 N 30 N 33 N 36 N 39 N 110 140 150 160 170 170 170 180 180 180 180 190 190 190 190 190 200 200 200 210 90 150 160 170 180 190 250 Sea Level (cm) Chlorophyll a (mg m -3 ) 160 E 165 E 170 E 175 E 180 175 W 170 W Figure 4. Geostrophic currents and sea surface height estimated from Topex/Poseidon satellite altimetry together with the track line and positions (dots) of the loggerhead turtle for (a) February 12–22, (b) March 13–23, and (c) April 12–22, 2001. The Kuroshio Extension Current is apparent as the strong eastward flow at 35 N. Figure 5. Geostrophic currents and sea surface height estimated from Topex/Poseidon satellite altimetry with loggerhead track line and positions (dots) for February 12–22, 2001 (top). Surface chlorophyll estimated from SeaWiFS ocean color together with loggerhead track line and positions (dots) for February 12–22, 2001 (bottom). Several olive ridley turtles used the equatorial current (Figs 6 and 7). One spent 40 days travelling on the NEC (Fig. 6). While in the NEC it traveled westward at an average speed of 71 cm s -1 (SE 12 cm s -1 , N = 73) compared with an average speed of 48 cm s -1 (SE 6 cm s -1 , N = 37) outside the NEC. The Topex/Poseiden estimates speed of the NEC during its 40 days in the NEC was 32 cm s -1 (SE 3 cm s -1 ). When the olive ridley was in the NEC it traveled at about 71 cm s -1 and as the NEC traveled at only 32 cm s -1 , the turtle appeared to be actively swimming and not just drifting as it rode the NEC. A second olive ridley used both the Equatorial Counter Current (ECC) and the NEC. It traveled eastward along the eastward-flowing ECC in September and then north and westward during October and November in the NEC (Fig. 7). 165 o E 168 o E 171 o E 174 o E 177 o E 180 177 o W 174 o W 8 o N 9 o N 10 o N 11 o N 12 o N 13 o N 14 o N 15 o N A. December 9 - 19 1997 165 o E 168 o E 171 o E 174 o E 177 o E 180 177 o W 174 o W 8 o N 9 o N 10 o N 11 o N 12 o N 13 o N 14 o N 15 o N 165 o E 168 o E 171 o E 174 o E 177 o E 180 177 o W 174 o W 8 o N 9 o N 10 o N 11 o N 12 o N 13 o N 14 o N 15 o N 165 o E 168 o E 171 o E 174 o E 177 o E 180 177 o W 174 o W 8 o N 9 o N 10 o N 11 o N 12 o N 13 o N 14 o N 15 o N B. December 19 - 29 1997 30 cm/sec 30 cm/sec C. December 29 1997 - January 8 1998 30 cm/sec D. January 8 - 18 1998 30 cm/sec In Press Polovina et al. 2003 Track for Olive Ridley 19584 over TOPEX Sea Surface Height Data 144 W 141 W 138 W 135 W 132 W 129 W 126 W 5N 7N 9N 11 N 13 N 15 N A. September 11 - 21 1997 30 cm/sec 144 W 141 W 138 W 135 W 132 W 129 W 126 W 5N 7N 9N 11 N 13 N 15 N B. October 11 - 21 1997 30 cm/sec 144 W 141 W 138 W 135 W 132 W 129 W 126 W 5N 7N 9N 11 N 13 N 15 N C. October 31 - November 10 1997 30 cm/sec Figure 6. Geostrophic current along the North Equatorial Current estimated from Topex/Poseidon satellite together with olive ridley track line and positions (dots) for: (a) December z, 1997, (b) December 19–29, 1997, (c) December 29, 1997–January 8, 1998, and (d) January 8–18, 1998. Figure 7. Geostrophic current along the North Equatorial Current estimated from Topex/Poseidon satellite together with olive ridley track line and positions (dots) for: (a) December 9–19, 1997, (b) December 19–29, 1997, (c) December 29, 1997–January 8, 1998, and (d) January 8–18, 1998. DISCUSSION Loggerhead and olive ridley turtles occupy habitats with different physical and biological characteristics and as a result utilize different types of forage and migration habitat. Loggerheads occupy the northern edge of the Subtropical Gyre and the Transition Zone, waters with a shallow thermocline, strong thermal and chlorophyll gradients, and surface convergent fronts. This results in relatively shallow dive depth distributions and most time spent at the surface, where they associate with eddies and fronts including the TZCF and the southern edge of the KEC. Olive ridleys are inhabit two different oceanic regions. Most remained in the center of the Subtropical Gyre characterized by warmer more vertically stratified water with a deeper thermocline without strong surface thermal and chlorophyll gradients. There they spend less time at the surface with a deeper depth distribution than loggerheads. Three olive ridleys occupied a second habitat consisting of major oceanic currents in the equatorial region, an area characterized by wind-induced upwelling and a shoaling of the thermocline, possibly allowing the olive ridleys to forage more shallowly in regions of surface convergences. Finally, our analyses show the value of combining satellite remote-sensed and ship-based physical and biological data with tracking data to further understand the oceanic habitat of pelagic animals and their movement patterns.

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Page 1: Forage and migration habitat of loggerhead (Caretta

Forage and migration habitat of loggerhead (Caretta caretta) and olive ridley (Lepidochelys olivacea) sea turtles in the central North Pacific OceanJeffrey J. Polovina1, George H. Balazs1, Evan A. Howell1, Denise M. Parker2, Michael P. Seki1 and Peter H. Dutton3

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Pacific Islands Fisheries Science Center, National Marine Fisheries Service, 2570 Dole Street, Honolulu, HI 96822-23962

Joint Institute for Marine and Atmospheric Research, University of Hawaii, 8604 La Jolla Drive, La Jolla, CA 920373

Southwest Fisheries Science Center, National Marine Fisheries Service, 8604 La Jolla Drive, La Jolla, CA 92037

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ABSTRACTSatellite telemetry from 26 loggerhead (Caretta caretta) and 10 olive ridley (Lepidochelys olivacea) sea turtles captured and released from pelagic longline fishing gear provided information on the turtles’ position and movement in the central North Pacific. These data together with environmental data from satellite remote sensing are used to describe the oceanic habitat used by these turtles. The results indicate that loggerheads travel westward, move seasonally north and south primarily through the region 28–40�N, and occupy sea surface temperatures (SST) of 15–25�C. Their dive depth distribution indicated that they spend 40% of their time at the surface and 90% of their time at depths <40 m. Loggerheads are found in association with fronts, eddies, and geostrophic currents. Specifically, the Transition Zone Chlorophyll Front (TZCF) and the southern edge of the Kuroshio Extension Current (KEC) appear to be important forage and migration habitats for loggerheads. In contrast, olive ridleys were found primarily south of loggerhead habitat in the region 8–31�N latitude, occupying warmer water with SSTs of 23–28�C. They have a deeper dive pattern than loggerheads, spending only 20% of their time at the surface and 60% shallower than 40 m. However, the three olive ridleys identified from genetics to be of western Pacific origin spent some time associated with major ocean currents, specifically the southern edge of the KEC, the North Equatorial Current (NEC), and the Equatorial Counter Current (ECC). These habitats were not used by any olive ridleys of eastern Pacific origin suggesting that olive ridleys from different populations may occupy different oceanic habitats.

150E 160E 170E 180 170W 160W 150W 140W 130W 120W5N

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Loggerhead

Olive Ridley

Figure 1. Track lines of 26 loggerhead (dashed) and 10 olive ridley (solid) turtles based on daily Argos reported positions from the period 1997–2001. Solid dots mark the start of each track line.

Telonics ST-10/ST-18 position transmitters were attached to 32 juvenile olive ridley and juvenile loggerhead Turtles from Hawaii-based longline fishing vessels from 1997-2001, with an additional 2 olive ridley and 2 loggerheads turtles instrumented with SDR-T10 dive recorders.

The spatial distribution of all track lines for the 36 turtles released with transmitters spanned an area from latitude 6–41�N and longitude 130�W–150�E (Fig. 1). Loggerheads were predominately north of about 28�N occupying the northern portion of the Subtropical Gyre and the Transition Zone, while the olive ridleys were generally south of 28�N in the Subtropical Gyre and the equatorial region (Fig. 1). All the loggerheads traveled westward with the exception of two on the eastern edge of the area that traveled southeast, while olive ridleys exhibited both eastward and westward movements.

Figure 2. Time-at-depth, dive depth, and daily maximum dive depth cumulative frequency distributions for all time periods combined: (a) both loggerheads combined, and (b) both olive ridleys combined.

From satellite SST estimated for each turtle position we estimated that olive ridleys primarily occupied a 5�C SST range, 23–28�C during all quarters, but within that range they appeared to be found centered at 24 and 27�C. Loggerheads occupied cooler water and covered a broader 10�C SST range, 15–25�C, occupying cooler temperatures in winter and spring and warmer temperature in the summer.

Both loggerheads exhibited relatively shallow dives with cumulative time-at-depth frequency distributions indicating that they spent 40% of their time at the surface and about 90% of their time at depths <40 m (Fig. 2). Both olive ridleys exhibited deeper diving than the loggerheads, with pooled cumulative time-at-depth frequency distributions indicating that they spent about 20% of their time at the surface and about 40% of their time deeper than 40 m (Fig. 2).

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Temperature-depth profiles indicated important differences between the thermal structure in the habitat used by the two turtle species. The olive ridley habitat was characterized by a deep thermocline at 100 m and minimal horizontal surface temperature fronts (Fig. 3). In contrast, the loggerhead habitat was characterized by a shallow thermocline at 50 m and several strong horizontal surface temperature fronts at 20 and 17�C located at 28 and 32�N, respectively (Fig. 3).

Similar depth differences in the chlorophyll-depth profiles were also observed, with the subsurface chlorophyll maximum above 100 m in the loggerhead habitat and reaching the surface at 33�N when it became the TZCF, a key feature for loggerhead habitat (Fig. 3). In the region of olive ridley habitat between 12 and 22�N latitude, the chlorophyll maximum was deeper than 100 m, although it shoaled to the surface at 10�N latitude at the northern edge of the NEC (Fig. 3).

Figure 3. Temperature depth and chlorophyll depth plots for (a) olive ridley habitat (10–22�N latitude, April 21–30, 2001), and (b) loggerhead habitat (25–34�N latitude, April 24–May 1, 1998).

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One interesting loggerhead track line shows the turtle’s westward trajectory just south of the strong eastward KEC, which is associated with the sharp gradient in SSH (Fig. 4). The turtle’s westward movement was often assisted by the westward flow on the north side of the counter-clockwise rotating eddies it encountered along 33�N (Fig. 4). During February–April 2001, the loggerhead position, as indicated by the black dots on the track line, was along the edge of a KEC meander. During February, March and April this meander developed into a cyclonic eddy, and the turtle occupied the edge of the eddy (Fig. 4). These meanders and eddies represent regions of enhanced surface chlorophyll as seen from surface chlorophyll estimated from SeaWiFS data for February 2001 (Fig. 5). The two cyclonic eddies at 165 and 180�E and the meander at 170�E seen in the altimetry data for February 12–22 are regions of enhanced surface chlorophyll (Fig. 5). The loggerhead remained for 3 months along the edge of the meander and eddy at 170�E which represents surface convergence zones and a likely site of concentrated prey for the loggerhead (Figs 4 and 5).

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Figure 4. Geostrophic currents and sea surface height estimated from Topex/Poseidon satellite altimetry together with the track line and positions (dots) of the loggerhead turtle for (a) February 12–22, (b) March 13–23, and (c) April 12–22, 2001. The Kuroshio Extension Current is apparent as the strong eastward flow at 35�N.

Figure 5. Geostrophic currents and sea surface height estimated from Topex/Poseidon satellite altimetry with loggerhead track line and positions (dots) for February 12–22, 2001 (top). Surface chlorophyll estimated from SeaWiFS ocean color together with loggerhead track line and positions (dots) for February 12–22, 2001 (bottom).

Several olive ridley turtles used the equatorial current (Figs 6 and 7). One spent 40 days travelling on the NEC (Fig. 6). While in the NEC it traveled westward at an average speed of 71 cm s-1 (SE 12 cm s-1, N = 73) compared with an average speed of 48 cm s-1 (SE 6 cm s-1, N = 37) outside the NEC. The Topex/Poseiden estimates speed of the NEC during its 40 days in the NEC was 32 cm s-1 (SE 3 cm s-1). When the olive ridley was in the NEC it traveled at about 71 cm s-1 and as the NEC traveled at only 32 cm s-1, the turtle appeared to be actively swimming and not just drifting as it rode the NEC. A second olive ridley used both the Equatorial Counter Current (ECC) and the NEC. It traveled eastward along the eastward-flowing ECC in September and then north and westward during October and November in the NEC (Fig. 7).

165oE 168oE 171oE 174oE 177oE 180 177oW 174oW8oN9oN

10oN11oN12oN13oN14oN15oN

A. December 9 - 19 1997

165oE 168oE 171oE 174oE 177oE 180 177oW 174oW8oN9oN

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165oE 168oE 171oE 174oE 177oE 180 177oW 174oW8oN9oN

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In Press Polovina et al. 2003

Track for Olive Ridley 19584 over TOPEX Sea Surface Height Data

144 W 141 W 138 W 135 W 132 W 129 W 126 W5 N

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Figure 6. Geostrophic current along the North Equatorial Current estimated from Topex/Poseidon satellite together with olive ridley track line and positions (dots) for: (a) December z, 1997, (b) December 19–29, 1997, (c) December 29, 1997–January 8, 1998, and (d) January 8–18, 1998.

Figure 7. Geostrophic current along the North Equatorial Current estimated from Topex/Poseidon satellite together with olive ridley track line and positions (dots) for: (a) December 9–19, 1997, (b) December 19–29, 1997, (c) December 29, 1997–January 8, 1998, and (d) January 8–18, 1998.

DISCUSSIONLoggerhead and olive ridley turtles occupy habitats with different physical and biological characteristics and as a result utilize different types of forage and migration habitat. Loggerheads occupy the northern edge of the Subtropical Gyre and the Transition Zone, waters with a shallow thermocline, strong thermal and chlorophyll gradients, and surface convergent fronts. This results in relatively shallow dive depth distributions and most time spent at the surface, where they associate with eddies and fronts including the TZCF and the southern edge of the KEC. Olive ridleys are inhabit two different oceanic regions. Most remained in the center of the Subtropical Gyre characterized by warmer more vertically stratified water with a deeper thermocline without strong surface thermal and chlorophyll gradients. There they spend less time at the surface with a deeper depth distribution than loggerheads. Three olive ridleys occupied a second habitat consisting of major oceanic currents in the equatorial region, an area characterized by wind-induced upwelling and a shoaling of the thermocline, possibly allowing the olive ridleys to forage more shallowly in regions of surface convergences. Finally, our analyses show the value of combining satellite remote-sensed and ship-based physical and biological data with tracking data to further understand the oceanic habitat of pelagic animals and their movement patterns.