Accuracy of ARGOS locations of pinnipeds at-sea estimated using Fastloc GPS

Background: ARGOS satellite telemetry is one of the most widely used methods to track the movements of free-ranging marine and terrestrial animals and is fundamental to studies of foraging ecology, migratory behavior and habitat-use. ARGOS location estimates do not include complete error estimations, and for many marine organisms, the most commonly acquired locations (Location Class 0, A, B, or Z) are provided with no declared error estimate.
Methodology/Principal Findings: We compared the accuracy of ARGOS locations to those obtained using Fastloc GPS from the same electronic tags on five species of pinnipeds: 9 California sea lions (Zalophus californianus), 4 Galapagos sea lions (Zalophus wollebaeki), 6 Cape fur seals (Arctocephalus pusillus pusillus), 3 Australian fur seals (A. p. doriferus) and 5 northern elephant seals (Mirounga angustirostris). These species encompass a range of marine habitats (highly pelagic vs coastal), diving behaviors (mean dive durations 2–21 min) and range of latitudes (equator to temperate). A total of 7,318 ARGOS positions and 27,046 GPS positions were collected. Of these, 1,105 ARGOS positions were obtained within five minutes of a GPS position and were used for comparison. The 68^th percentile ARGOS location errors as measured in this study were LC-3
0.49 km, LC-2 1.01 km, LC-1 1.20 km, LC-0 4.18 km, LC-A 6.19 km, LC-B 10.28 km.
Conclusions/Significance: The ARGOS errors measured here are greater than those provided by ARGOS, but within the range of other studies. The error was non-normally distributed with each LC highly right-skewed. Locations of species that make short duration dives and spend extended periods on the surface (sea lions and fur seals) had less error than species like elephant seals that spend more time underwater and have shorter surface intervals. Supplemental data (S1) are provided allowing the creation of density distributions that can be used in a variety of filtering algorithms to improve the quality of ARGOS tracking data.

Factors influencing the winter haulout behaviour of Weddell seals : consequences for satellite telemetry

An understanding of pinniped haulout behaviour can provide information on how animals interact with their environment, their foraging strategies and the behaviour of their prey. For Weddell seals Leptonychotes weddelli, knowledge of haulout patterns and the environmental drivers of haulout is comprehensive for the austral spring and summer, but is poorly described outside this period. Pinniped behaviour is commonly examined using satellite telemetry; however, it is unclear whether the behavioural state of the seal can itself influence data acquisition. We examined haulout behaviour of female Weddell seals in eastern Antarctica over 3 winters (March to August, 2006–2008) using satellite-linked data loggers. Haulout behaviour followed a diel cycle with predominantly nocturnal haulouts. The environmental variables wind speed and temperature were influential on haulout, with seals tending to haul out more in lower winds and at higher temperatures. Haulout duration decreased across the winter, as did the number of satellite uplinks, suggesting that haulout duration played a role in data acquisition. There was no evidence of a decline in tag condition over this period, indicating that the decrease in uplinks was primarily the result of the winter behaviour of the seals. Overall, the number of haulout locations in the data set was over-represented, with more uplinks per hour occurring when the seals were hauled out than when they were at sea. For Weddell seals and other ice seals, tracking data that include haulout locations can bias spatial representations of foraging behaviour (e.g. haulout sites may be mistaken for regions of area-restricted search).

Conservation hotspots : implications of intense spatial area use by breeding male and female loggerheads at the Mediterranean’s largest rookery

The implementation of appropriate protection measures for endangered species in protected areas requires knowledge of their fine-scale habitat use. In May and June of 2006 and 2007, we used GPS loggers (some linked to the Argos system) and a conventional Argos transmitter to track male and female loggerhead turtles Caretta caretta in the vicinity of the breeding area of Laganas Bay within the National Marine Park of Zakynthos, Greece. We obtained (1) 9681 useable locations (mean: 1383 locations ind.–1; range: 519 to 2198 locations) from Tracktag GPS loggers attached to 7 females for a mean duration of 34 d (range: 17 to 52 d); (2) 1245 useable locations (mean: 311 locations ind.–1; range: 38 to 1110 locations) from 4 males fitted with Fastloc Argos tags for a mean duration of 29 d (range: 3 to 51 d) and (3) 100 locations from 1 male fitted with a conventional Argos satellite tag tracked for 128 d. GPS data indicated that before the onset of nesting, both males and females primarily used an area within 500 m of the shore along a core 9 km stretch of coastline, where existing protective legislation requires strengthening. Our observations suggest that a 76.7% female-biased operational sex ratio, measured previously from in-water surveys, may represent a realistic sex ratio estimate in the period before nesting starts. In the first month following the onset of nesting, female spatial distribution remained similar, whereas most males departed for distant areas presumably to forage. Our study provides quantitative evidence of the need to improve the management planning and conservation measures to protect sea turtles in a coastal breeding area, and new insights on male turtle migration.

A biologist's guide to assessing ocean currents : a review

We review how ocean currents are measured (in both Eulerian and Lagrangian frameworks), how they are inferred from satellite observations, and how they are simulated in ocean general circulation models (OGCMs). We then consider the value of these ‘direct’ (in situ) and ‘indirect’ (inferred, simulated) approaches to biologists investigating current-induced drift of strong-swimming vertebrates as well as dispersion of small organisms in the open ocean. We subsequently describe 2 case studies. In the first, OGCM-simulated currents were compared with satellite-derived currents; analyses suggest that the 2 methods yield similar results, but that each has its own limitations and associated uncertainty. In the second analysis, numerical methods were tested using Lagrangian drifter buoys. Results indicated that currents simulated in OGCMs do not capture all details of buoy trajectories, but do successfully resolve most general aspects of current flows. We thus recommend that the errors and uncertainties in ocean current measurements, as well as limitations in spatial and temporal resolution of the surface current data, need to be considered in tracking studies that incorporate oceanographic data. Whenever possible, cross-validation of the different methods (e.g. indirect estimates versus buoy trajectories) should be undertaken before a decision is reached about which technique is best for a specific application.

Movement patterns of green turtles in Brazilian coastal waters described by satellite tracking and flipper tagging

The movements of 8 green turtles Chelonia mydas in Brazilian coastal waters were tracked using transmitters linked to the Argos system for periods of between 1 and 197 d. These were the first tracking data gathered on juveniles of this species in this important foraging ground. Information was integrated with that collected over a decade using traditional flipper-tagging methods at the same site. Both satellite telemetry and flipper tagging suggested that turtles undertook 1 of 3 general patterns of behaviour: pronounced long range movements (>100 km), moderate range movements (<100 km) or extended residence very close to the capture/release site. There seemed to be a general tendency for the turtles recaptured/tracked further afield to have been among the larger turtles captured. Satellite tracking of 5 turtles which moved from the release site showed that they moved through coastal waters; a factor which is likely to predispose migrating turtles to incidental capture as a result of the prevailing fishing methods in the region. The movements of the 3 turtles who travelled less than 100 km from the release site challenge previous ideas relating to home range in green turtles feeding in sea grass pastures. We hypothesise that there may be a fundamental difference in the pattern of habitat utilisation by larger green turtles depending on whether they are feeding on seagrass or macroalgae. Extended tracking of 2 small turtles which stayed near the release point showed that small juvenile turtles, whilst in residence in a particular feeding ground, can also exhibit high levels of site-fidelity with home ranges of the order of several square kilometers.

Movement patterns for a critically endangered species, the Leatherback Turtle (Dermochelys coriacea), linked to foraging success and population status

Foraging success for pelagic vertebrates may be revealed by horizontal and vertical movement patterns. We show markedly different patterns for leatherback turtles in the North Atlantic versus Eastern Pacific, which feed on gelatinous zooplankton that are only occasionally found in high densities. In the Atlantic, travel speed was characterized by two modes, indicative of high foraging success at low speeds (<15 km d−1) and transit at high speeds (20–45 km d−1). Only a single mode was evident in the Pacific, which occurred at speeds of 21 km d−1 indicative of transit. The mean dive depth was more variable in relation to latitude but closer to the mean annual depth of the thermocline and nutricline for North Atlantic than Eastern Pacific turtles. The most parsimonious explanation for these findings is that Eastern Pacific turtles rarely achieve high foraging success. This is the first support for foraging behaviour differences between populations of this critically endangered species and suggests that longer periods searching for prey may be hindering population recovery in the Pacific while aiding population maintenance in the Atlantic.

Satellite telemetry suggests high levels of fishing-induced mortality in marine turtles

Long-term records of nesting numbers, or proxies to nesting numbers, show a precipitous decline in the size of many sea turtle populations. Population declines are most frequently attributed to fisheries bycatch, although direct quantification of this level of mortality is rare. We used satellite-tracking records for turtles in the Mediterranean Sea and Pacific, Atlantic and Indian Oceans to identify when turtles had been captured. Evidence for capture came from a combination of an increase in good quality locations from transmitters, transmitters moving inland to coastal towns and villages, and on-board submergence data, showing that transmitters had come out of the water. A high level of mortality was calculated, confirming current concerns regarding the outlook for sea turtles.

Long-term satellite telemetry of the movements and habitat utilisation by green turtles in the Mediterranean

There is a relative paucity of data regarding the at-sea distribution and behaviour of marine turtles. This is especially true for the critically endangered green turtle Chelonia mydas population in the Mediterranean. Six adult female green turtles were equipped with satellite transmitters and tracked for periods of between 28 and 293 d following their final nesting of the season in northern Cyprus. Data elucidated hitherto unknown migratory pathways and highlighted the importance of North African coastal waters as feeding habitat for adults of this species. For three individuals, instruments transmitted detailed information on dive depth, dive duration and water temperature which afforded novel insights into behaviour during different stages of migration, feeding in the foraging grounds and most remarkably, during a period of midwinter diapause when water temperatures were generally below 25°C. Turtles showed fidelity to specific shallow inshore feeding areas and moved offshore to deeper wintering sites.

The diving behaviour of green turtles undertaking oceanic migration to and from Ascension Island: dive durations, dive profiles and depth distribution

Satellite telemetry was used to record the submergence duration of green turtles (Chelonia mydas) as they migrated from Ascension Island to Brazil (N=12 individuals) while time/depth recorders (TDRs) were used to examine the depth distribution and dive profiles of individuals returning to Ascension Island to nest after experimental displacement (N=5 individuals). Satellite telemetry revealed that most submergences were short (<5 min) but that some submergences were longer (>20 min), particularly at night. TDRs revealed that much of the time was spent conducting short (2–4 min), shallow (approximately 0.9–1.5 m) dives, consistent with predictions for optimisation of near-surface travelling, while long (typically 20–30 min), deep (typically 10–20 m) dives had a distinctive profile found in other marine reptiles. These results suggest that green turtles crossing the Atlantic do not behave invariantly, but instead alternate between periods of travelling just beneath the surface and diving deeper. These deep dives may have evolved to reduce silhouetting against the surface, which would make turtles more susceptible to visual predators such as large sharks.

Changes in behaviour during the inter-nesting period and post-nesting migration for Ascension Island green turtles

Satellite transmitters were attached to green turtles Chelonia mydas while they were nesting on Ascension Island in the South Atlantic (7°57'S, 14°22'W) and individuals were subsequently monitored during the inter-nesting period and the post-nesting migration to Brazil. During the inter-nesting period, data from the transmitters suggested that turtles generally stayed within 5 km of the nesting beach on which they had originally been observed. During both the inter-nesting period and migration, turtles were submerged the vast majority (>95%) of the time, suggesting that they neither basked at the surface nor drifted passively during migration to any great extent. There was a clear dichotomy in submergence behaviour, with submergences tending to be of short duration during post-nesting migration (mean = 7.3 min, 3318 h of data from 5 individuals) and of longer duration during the inter-nesting period (mean = 22.1 min, 714 h of data from 5 different individuals). As submergence duration is generally linked to activity levels in sea turtles, this pattern suggests that turtles were comparatively inactive during the inter-nesting period and comparatively active during migration. During both the inter-nesting period and the post-nesting migration, diel submergence patterns were detected with dive duration tending to be longer at night. As the turtles migrated WSW from Ascension Island, there was a reduction in their speed of travel. A numerical model of the near-surface currents suggested that this reduction was associated with the weakening of the WSW flow of the prevailing South Atlantic Equatorial Current.

When surfacers do not dive : multiple significance of extended surface times in marine turtles

Marine turtles spend more than 90% of their life underwater and have been termed surfacers as opposed to divers. Nonetheless turtles have been reported occasionally to float motionless at the surface but the reasons for this behaviour are not clear. We investigated the location, timing and duration of extended surface times (ESTs) in 10 free-ranging loggerhead turtles (Caretta caretta) and the possible relationship to water temperature and diving activity recorded via satellite relay data loggers for 101–450 days. For one turtle that dived only in offshore areas, ESTs contributed 12% of the time whereas for the other turtles ESTs contributed 0.4–1.8% of the time. ESTs lasted on average 90 min but were mostly infrequent and irregular, excluding the involvement of a fundamental regulatory function. However, 82% of the ESTs occurred during daylight, mostly around noon, suggesting a dependence on solar radiation. For three turtles, there was an appreciable (7°C to 10.5°C) temperature decrease with depth for dives during periods when ESTs occurred frequently, suggesting a re-warming function of EST to compensate for decreased body temperatures, possibly to enhance digestive efficiency. A positive correlation between body mass and EST duration supported this explanation. By contrast, night-active turtles that exceeded their calculated aerobic dive limits in 7.6–16% of the dives engaged in nocturnal ESTs, probably for lactate clearance. This is the first evidence that loggerhead turtles may refrain from diving for at least two reasons, either to absorb solar radiation or to recover from anaerobic activity.

Leatherback turtles satellite-tagged in European waters

The North Atlantic is considered a stronghold for the critically endangered leatherback sea turtle. However, limited information exists regarding the movements of individuals to and from the seas off Europe’s northwesterly fringe, an area where leatherbacks have been historically sighted for the past 200 yr. Here, we used satellite telemetry to record the movements and behaviour of 2 individuals bycaught in fisheries off the southwest coast of Ireland. The turtle T1 (tagged 1 September 2005; female; tracked 375 d) immediately travelled south via Madeira and the Canaries, before residing in West African waters for 3 mo. In spring, T1 migrated north towards Newfoundland where transmissions ceased. T2 (29 June 2006; male; 233 d) travelled south for a short period before spending 66 d west of the Bay of Biscay, an area previously asserted as a high-use area for leatherbacks. This prolonged high latitude summer residence corresponded with a mesoscale feature evident from satellite imagery, with the implication that this turtle had found a rich feeding site. A marked change in dive behaviour was apparent as the turtle exited this feature and provided useful insights on leatherback diving behaviour. T2 headed south in October 2006, and performed the deepest-ever dive recorded by a reptile (1280 m) southwest of Cape Verde. Unlike T1, T2 swam southwest towards Brazil before approaching the major nesting beaches of French Guiana and Surinam. Importantly, these tracks document the movement of leatherbacks from one of the remotest foraging grounds in the North Atlantic.