Jean-Yves Georges

Marine animal behaviour: neglecting ocean currents can lead us up the wrong track

Tracks of marine animals in the wild, now increasingly acquired by electronic tagging of individuals, are of prime interest not only to identify habitats and high-risk areas, but also to gain detailed information about the behaviour of these animals. Using recent satellite-derived current estimates and leatherback turtle (Dermochelys coriacea) tracking data, we demonstrate that oceanic currents, usually neglected when analysing tracking data, can substantially distort the observed trajectories. Consequently, this will affect several important results deduced from the analysis of tracking data, such as the evaluation of the orientation skills and the energy budget of animals or the identification of foraging areas. We conclude that currents should be systematically taken into account to ensure the unbiased interpretation of tracking data, which now play a major role in marine conservation biology.

MATERNAL CARE IN THE SUBANTARCTIC FUR SEALS ON AMSTERDAM ISLAND

This paper reports the first study of maternal input and care from birth to weaning in a fur seal with a long pup-rearing period: the subantarctic fur seal Arctocephalus tropicalis breeding on the temperate Amsterdam Island, Indian Ocean. The protracted wean- ing period provided the opportunity for examination of maternal care in relation to seasonal changes in the requirements of the mother-pup pair and environmental conditions. During the reproductive season 1995-1996, maternal care was investigated in terms of provisioning (maternal attendance) pattern while diving effort was investigated using time depth recorders in summer and winter. Maternal input was calculated in terms of the absolute rate of pup mass gain and, ultimately, pup growth rate and pup body mass at weaning. Lactating subantarctic fur seals perform one of the longest attendance cycles described in fur seals, spending on average 11-23 d at sea from summer to winter. The time mothers spend ashore suckling their pup is also long (;4 d) but remains constant throughout the year. Throughout the year, maternal input should be described as follows: mothers spending a long time at sea store a large amount of body reserves that provide them a good body condition. Con- sequently, they spend a long time ashore to transfer their body reserves to their pups. However, mothers spending short attendance periods increase the mass transfer efficiency, probably by decreasing their metabolic overhead. In summer, maternal care was mostly controlled by pup traits: maternal absences appeared to be controlled by pup fasting ability, while maternal input was controlled by pup ingestion ability, i.e., pup body size and the time the pup was suckling. In fall, pups were no longer limited in milk ingestion, and maternal input was mostly controlled by maternal traits (e.g., body length and experience). In winter, maternal input decreased as the pup became older despite an increase in maternal diving effort. We propose that, in winter, maternal requirements increase, probably in response to increasing costs of gestation and because of a decrease in food resource avail- ability. Pups whose mother performed short and regular foraging trips grew faster and were heavier at weaning than other pups. This is discussed in term of pup fasting endurance and maternal experience. Finally, we found a window of foraging trip durations that maximizes the net rate of energy acquisition of the pup, suggesting that in subantarctic fur seals there may not exist one optimal maternal attendance pattern, but a range of patterns promoting the same maternal fitness.

How long does a dive last? Foraging decisions by breath-hold divers in a patchy environment: a test of a simple model

*Sea Mammal Research Unit, University of St AndrewsyCentre National de la Recherche Scientifique, Institut Pluridisciplinaire Hubert Curien, Departement Ecologie,Physiologie et Ethologie (DEPE)(Received 1 March 2006; initial acceptance 25 April 2006;final acceptance 26 June 2006; published online 10 July 2007; MS. number: 8866R)Many theoretical models have been proposed to explain and predict the behaviour of air-breathing diversexploiting a food resource underwater. Many field observations of the behaviour of divers do not fit withthe prediction that to maximize energetic gain divers should dive close to their aerobic diving limits. In anattempt to explain this paradox, Thompson & Fedak (2001, Animal Behaviour, 61, 286e297) proposeda model of diving behaviour that takes into account patchily distributed prey patches of varying quality.We tested this model experimentally in a simulated foraging set-up. We measured the diving behaviour ofgrey seals, Halichoerus grypus, diving to patches of varying prey density and distance from the surface. Ourresults were equivocal with respect to the model predictions. Seals responded to prey density, leaving low-quality patches earlier. However, this pattern was still evident at long dive distances, contrary to theprediction that during deep dives seals should stay at a patch regardless of prey density. While seals max-imized dive durations at high prey densities and long distances, they did not do so at short distances. Theapparent quitting strategy of the seals always produced higher net rates of energy gain than would havebeen achieved if they had remained at the foraging site up to their aerobic dive limit on every dive. Theseresults indicate that seals’ diving behaviour, particularly bottom duration, may indicate the relative preyavailability in their environment.

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.

Energy expenditure of freely swimming adult green turtles ( Chelonia mydas ) and its link with body acceleration

SUMMARY Marine turtles are globally threatened. Crucial for the conservation of these large ectotherms is a detailed knowledge of their energy relationships, especially their at-sea metabolic rates, which will ultimately define population structure and size. Measuring metabolic rates in free-ranging aquatic animals, however, remains a challenge. Hence, it is not surprising that for most marine turtle species we know little about the energetic requirements of adults at sea. Recently, accelerometry has emerged as a promising tool for estimating activity-specific metabolic rates of animals in the field. Accelerometry allows quantification of the movement of animals (ODBA/PDBA, overall/partial dynamic body acceleration), which, after calibration, might serve as a proxy for metabolic rate. We measured oxygen consumption rates () of adult green turtles (Chelonia mydas; 142.1±26.9 kg) at rest and when swimming within a 13 m-long swim channel, using flow-through respirometry. We investigated the effect of water temperature (Tw) on turtle and tested the hypothesis that turtle body acceleration can be used as a proxy for . Mean mass-specific () of six turtles when resting at a Tw of 25.8±1.0°C was 0.50±0.09 ml min–1 kg–0.83. increased significantly with Tw and activity level. Changes in were paralleled by changes in respiratory frequency (fR). Deploying bi-axial accelerometers in conjunction with respirometry, we found a significant positive relationship between and PDBA that was modified by Tw. The resulting predictive equation was highly significant (r2=0.83, P<0.0001) and associated error estimates were small (mean algebraic error 3.3%), indicating that body acceleration is a good predictor of in green turtles. Our results suggest that accelerometry is a suitable method to investigate marine turtle energetics at sea.

Atlantic Leatherback Migratory Paths and Temporary Residence Areas

Background Sea turtles are long-distance migrants with considerable behavioural plasticity in terms of migratory patterns, habitat use and foraging sites within and among populations. However, for the most widely migrating turtle, the leatherback turtle Dermochelys coriacea, studies combining data from individuals of different populations are uncommon. Such studies are however critical to better understand intra- and inter-population variability and take it into account in the implementation of conservation strategies of this critically endangered species. Here, we investigated the movements and diving behaviour of 16 Atlantic leatherback turtles from three different nesting sites and one foraging site during their post-breeding migration to assess the potential determinants of intra- and inter-population variability in migratory patterns. Methodology/Principal Findings Using satellite-derived behavioural and oceanographic data, we show that turtles used Temporary Residence Areas (TRAs) distributed all around the Atlantic Ocean: 9 in the neritic domain and 13 in the oceanic domain. These TRAs did not share a common oceanographic determinant but on the contrary were associated with mesoscale surface oceanographic features of different types (i.e., altimetric features and/or surface chlorophyll a concentration). Conversely, turtles exhibited relatively similar horizontal and vertical behaviours when in TRAs (i.e., slow swimming velocity/sinuous path/shallow dives) suggesting foraging activity in these productive regions. Migratory paths and TRAs distribution showed interesting similarities with the trajectories of passive satellite-tracked drifters, suggesting that the general dispersion pattern of adults from the nesting sites may reflect the extent of passive dispersion initially experienced by hatchlings. Conclusions/Significance Intra- and inter-population behavioural variability may therefore be linked with initial hatchling drift scenarios and be highly influenced by environmental conditions. This high degree of behavioural plasticity in Atlantic leatherback turtles makes species-targeted conservation strategies challenging and stresses the need for a larger dataset (>100 individuals) for providing general recommendations in terms of conservation.

Milking strategy in subantarctic fur seals Arctocephalus tropicalis breeding on Amsterdam Island: evidence from changes in milk composition.

Milk composition was investigated throughout the 10‐mo pup‐rearing period in subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island. The mean milk composition was \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm} \usepackage{mathrsfs} \usepackage{pifont} \usepackage{stmaryrd} \usepackage{textcomp} \usepackage{portland,xspace} \usepackage{amsmath,amsxtra} \usepackage[OT2,OT1]{fontenc} \newcommand\cyr{ \renewcommand\rmdefault{wncyr} \renewcommand\sfdefault{wncyss} \renewcommand\encodingdefault{OT2} \normalfont \selectfont} \DeclareTextFontCommand{\textcyr}{\cyr} \pagestyle{empty} \DeclareMathSizes{10}{9}{7}{6} \begin{document} \landscape

Pleated turtle escapes the box – shape changes in Dermochelys coriacea

42.8\% \pm 5.7\%