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It is usually beneficial for species to restrict activity to a particular phase of the 24-hour cycle as this enables the development of morphological and behavioural adaptations to enhance survival under specific biotic and abiotic conditions. Sloth activity patterns are thought to be strongly related to the environmental conditions due to the metabolic consequences of having a low and highly variable core body temperature. Understanding the drivers of sloth activity and their ability to withstand environmental fluctuations is of growing importance for the development of effective conservation measures, particularly when we consider the vulnerability of tropical ecosystems to climate change and the escalating impacts of anthropogenic activities in South and Central America. Unfortunately, the cryptic nature of sloths makes long term observational research difficult and so there is very little existing literature examining the behavioural ecology of wild sloths. Here, we used micro data loggers to continuously record, for the first time, the behaviour of both Bradypus and Choloepus sloths over periods of days to weeks. We investigate how fluctuations in the environmental conditions affect the activity of sloths inhabiting a lowland rainforest on the Caribbean coast of Costa Rica and examined how this might relate to their low power lifestyle. Both Bradypus and Choloepus sloths were found to be cathemeral in their activity, with high levels of between-individual and within-individual variation in the amounts of time spent active, and in the temporal distribution of activity over the 24-hour cycle. Daily temperature did not affect activity, although Bradypus sloths were found to show increased nocturnal activity on colder nights, and on nights following colder days. Our results demonstrate a distinct lack of synchronicity within the same population, and we suggest that this pattern provides sloths with the flexibility to exploit favourable environmental conditions whilst reducing the threat of predation.

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Understanding how animals move in dense environments where vision is compromised is a major challenge. We used GPS and dead-reckoning to examine the movement of Magellanic penguins commuting through vegetation that precluded long-distance vision. Birds leaving the nest followed the shortest, quickest route to the sea (the 'ideal path', or 'I-path') but return tracks depended where the birds left the water. Penguins arriving at the beach departure spot mirrored the departure. Most of those landing at a distance from the departure spot travelled slowly, obliquely to the coast at a more acute angle than a beeline trajectory to the nest. On crossing their I-path, these birds then followed this route quickly to their nests. This movement strategy saves birds distance, time and energy compared to a route along the beach and the into the colony on the I-track and saves time and energy compared to a beeline trajectory which necessitates slow travel in unfamiliar areas. This suggests that some animals adopt tactics that take them to an area where their navigational capacities are enhanced for efficient travel in challenging environments.

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Sloths are notoriously slow and consequently have limited dispersal ability, which makes them particularly vulnerable to the effects of habitat fragmentation and degradation. Sloths in Costa Rica are considered of conservation concern due to habitat loss, livestock production and increasing urbanization. Reintroductions from rescue centres are commonplace across the country, yet their genetic diversity and population structure are unknown, and there is currently little consideration of the genetic background prior to intervention or releases. We used microsatellite analysis to undertake the first exploratory investigation into sloth population genetics in Costa Rica. Using data from 98 two-fingered sloths (Choloepus hoffmanni) from four different geographic regions, we determined the presence of four potential genetic groups, three of them with minimal population structuring despite the limited dispersal ability and presence of physical barriers. Sloths from the North appear to represent a highly distinct population that we propose may require management as a discrete unit for conservation. We stress the need for additional analyses to better understand the genetic structure and diversity of North andWest regions and suggest that rescue facilities in Costa Rica should consider the genetic background of rehabilitated sloths when planning future reintroductions. Our results also highlight the threat posed by physical isolation due to widespread urbanization and agriculture expansion for a species with a weak dispersal ability.

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BACKGROUND: Precision Livestock Farming (PLF) is a promising approach to minimize the conflicts between socio-economic activities and landscape conservation. However, its application on extensive systems of livestock production can be challenging. The main difficulties arise because animals graze on large natural pastures where they are exposed to competition with wild herbivores for heterogeneous and scarce resources, predation risk, adverse weather, and complex topography. Considering that the 91% of the world's surface devoted to livestock production is composed of extensive systems (i.e., rangelands), our general aim was to develop a PLF methodology that quantifies: (i) detailed behavioural patterns, (ii) feeding rate, and (iii) costs associated with different behaviours and landscape traits. METHODS: For this, we used Merino sheep in Patagonian rangelands as a case study. We combined data from an animal-attached multi-sensor tag (tri-axial acceleration, tri-axial magnetometry, temperature sensor and Global Positioning System) with landscape layers from a Geographical Information System to acquire data. Then, we used high accuracy decision trees, dead reckoning methods and spatial data processing techniques to show how this combination of tools could be used to assess energy balance, predation risk and competition experienced by livestock through time and space. RESULTS: The combination of methods proposed here are a useful tool to assess livestock behaviour and the different factors that influence extensive livestock production, such as topography, environmental temperature, predation risk and competition for heterogeneous resources. We were able to quantify feeding rate continuously through time and space with high accuracy and show how it could be used to estimate animal production and the intensity of grazing on the landscape. We also assessed the effects of resource heterogeneity (inferred through search times), and the potential costs associated with predation risk, competition, thermoregulation and movement on complex topography. DISCUSSION: The quantification of feeding rate and behavioural costs provided by our approach could be used to estimate energy balance and to predict individual growth, survival and reproduction. Finally, we discussed how the information provided by this combination of methods can be used to develop wildlife-friendly strategies that also maximize animal welfare, quality and environmental sustainability.

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RATIONALE: Food provisioning is considered one of the main traits affecting offspring fitness. Differences in food provisioning between sexes, particularly in dimorphic species, could affect the amount and type of food provided, due to differences in the amount of food carried to the nest as a result of differential resources exploitation. Quantitative evidence for sexual differences in food provisioning by parents in penguins is scarce. The Magellanic penguin is moderately sexually dimorphic and breeds along a broad latitudinal range, with birds north and south of this range being essentially dietary specialists while those at intermediate latitudes consume a more diverse diet. METHODS: We used stable isotope analysis of carbon and nitrogen by isotope ratio mass spectrometry to examine if there was a differential parental contribution to chicks in ten Magellanic penguin colonies throughout its latitudinal breeding distribution. We used the heuristic Euclidean isotopic distance (ED) and individual isotope distances between the chicks and their parents as a proxy for diet similarity (the smaller the distance, the more similar the diet). RESULTS: The analysis showed that chicks tended to have a more similar diet to that of their male parent and that this pattern was more evident at colonies and in seasons where penguins had a more diverse diet, which could be explained by differences in diet between parents. Distance in δ15 N values, but not in δ13 C values, differed between both sexes and their chicks in all the pairs sampled, suggesting that δ15 N values drive the differences found in ED between chicks and their parents. CONCLUSIONS: We have developed an approach that provides the first assessment of the extent of differential food provisioning between male and female Magellanic penguins. Results suggest chicks have a diet more similar to that of their male parent, probably related to the higher trophic level of male penguin prey.

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Penguins are major consumers in the southern oceans although quantification of this has been problematic. One suggestion proposes the use of points of inflection in diving profiles ('wiggles') for this, a method that has been validated for the estimation of prey consumption by Magellanic penguins (Spheniscus magellanicus) by Simeone and Wilson (2003). Following them, we used wiggles from 31 depth logger-equipped Magellanic penguins foraging from four Patagonian colonies; Punta Norte (PN), Bahía Bustamente (BB), Puerto Deseado (PD) and Puerto San Julián (PSJ), all located in Argentina between 42-49° S, to estimate the prey captured and calculate the catch per unit time (CPUT) for birds foraging during the early chick-rearing period. Numbers of prey caught and CPUT were significantly different between colonies. Birds from PD caught the highest number of prey per foraging trip, with CPUT values of 68±19 prey per hour underwater (almost two times greater than for the three remaining colonies). We modeled consumption from these data and calculate that the world Magellanic penguin population consumes about 2 million tons of prey per year. Possible errors in this calculation are discussed. Despite this, the analysis of wiggles seems a powerful and simple tool to begin to quantify prey consumption by Magellanic penguins, allowing comparison between different breeding sites. The total number of wiggles and/or CPUT do not reflect, by themselves, the availability of food for each colony, as the number of prey consumed by foraging trip is strongly associated with the energy content and wet mass of each colony-specific 'prey type'. Individuals consuming more profitable prey could be optimizing the time spent underwater, thereby optimizing the energy expenditure associated with the dives.

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Dynamic body acceleration (DBA) has been used as a proxy for energy expenditure in logger-equipped animals, with researchers summing the acceleration (overall dynamic body acceleration--ODBA) from the three orthogonal axes of devices. The vector of the dynamic body acceleration (VeDBA) may be a better proxy so this study compared ODBA and VeDBA as proxies for rate of oxygen consumption using humans and 6 other species. Twenty-one humans on a treadmill ran at different speeds while equipped with two loggers, one in a straight orientation and the other skewed, while rate of oxygen consumption (VO2) was recorded. Similar data were obtained from animals but using only one (straight) logger. In humans, both ODBA and VeDBA were good proxies for VO2 with all r(2) values exceeding 0.88, although ODBA accounted for slightly but significantly more of the variation in VO2 than did VeDBA (P<0.03). There were no significant differences between ODBA and VeDBA in terms of the change in VO2 estimated by the acceleration data in a simulated situation of the logger being mounted straight but then becoming skewed (P = 0.744). In the animal study, ODBA and VeDBA were again good proxies for VO2 with all r(2) values exceeding 0.70 although, again, ODBA accounted for slightly, but significantly, more of the variation in VO2 than did VeDBA (P<0.03). The simultaneous contraction of muscles, inserted variously for limb stability, may produce muscle oxygen use that at least partially equates with summing components to derive DBA. Thus, a vectorial summation to derive DBA cannot be assumed to be the more 'correct' calculation. However, although within the limitations of our simple study, ODBA appears a marginally better proxy for VO2. In the unusual situation where researchers are unable to guarantee at least reasonably consistent device orientation, they should use VeDBA as a proxy for VO2.

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Variation in the physical characteristics of the environment should impact the movement energetics of animals. Although cognizance of this may help interpret movement ecology, determination of the landscape-dependent energy expenditure of wild animals is problematic. We used accelerometers in animal-attached tags to derive energy expenditure in 54 free-living imperial cormorants Phalacrocorax atriceps and construct an energy landscape of the area around a breeding colony. Examination of the space use of a further 74 birds over 4 years showed that foraging areas selected varied considerably in distance from the colony and water depth, but were characterized by minimal power requirements compared with other areas in the available landscape. This accords with classic optimal foraging concepts, which state that animals should maximize net energy gain by minimizing costs where possible and show how deriving energy landscapes can help understand how and why animals distribute themselves in space.

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Locomotion is one of the major energetic costs faced by animals and various strategies have evolved to reduce its cost. Birds use interspersed periods of flapping and gliding to reduce the mechanical requirements of level flight while undergoing cyclical changes in flight altitude, known as undulating flight. Here we equipped free-ranging marine vertebrates with accelerometers and demonstrate that gait patterns resembling undulating flight occur in four marine vertebrate species comprising sharks and pinnipeds. Both sharks and pinnipeds display intermittent gliding interspersed with powered locomotion. We suggest, that the convergent use of similar gait patterns by distinct groups of animals points to universal physical and physiological principles that operate beyond taxonomic limits and shape common solutions to increase energetic efficiency. Energetically expensive large-scale migrations performed by many vertebrates provide common selection pressure for efficient locomotion, with potential for the convergence of locomotory strategies by a wide variety of species.

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Animals respond to environmental variation by exhibiting a number of different behaviours and/or rates of activity, which result in corresponding variation in energy expenditure. Successful animals generally maximize efficiency or rate of energy gain through foraging. Quantification of all features that modulate energy expenditure can theoretically be modelled as an animal energetic niche or power envelope; with total power being represented by the vertical axis and n-dimensional horizontal axes representing extents of processes that affect energy expenditure. Such an energetic niche could be used to assess the energetic consequences of animals adopting particular behaviours under various environmental conditions. This value of this approach was tested by constructing a simple mechanistic energetics model based on data collected from recording devices deployed on 41 free-living Magellanic penguins (Spheniscus magellanicus), foraging from four different colonies in Argentina and consequently catching four different types of prey. Energy expenditure was calculated as a function of total distance swum underwater (horizontal axis 1) and maximum depth reached (horizontal axis 2). The resultant power envelope was invariant, irrespective of colony location, but penguins from the different colonies tended to use different areas of the envelope. The different colony solutions appeared to represent particular behavioural options for exploiting the available prey and demonstrate how penguins respond to environmental circumstance (prey distribution), the energetic consequences that this has for them, and how this affects the balance of energy acquisition through foraging and expenditure strategy.

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In the face of the physical and physiological challenges of performing breath-hold deep dives, marine vertebrates have evolved different strategies. Although behavioural strategies in marine mammals and seabirds have been investigated in detail, little is known about the deepest-diving reptile - the leatherback turtle (Dermochelys coriacea). Here, we deployed tri-axial accelerometers on female leatherbacks nesting on St Croix, US Virgin Islands, to explore their diving strategy. Our results show a consistent behavioural pattern within dives among individuals, with an initial period of active swimming at relatively steep descent angles (∼-40 deg), with a stroke frequency of 0.32 Hz, followed by a gliding phase. The depth at which the gliding phase began increased with the maximum depth of the dives. In addition, descent body angles and vertical velocities were higher during deeper dives. Leatherbacks might thus regulate their inspired air-volume according to the intended dive depth, similar to hard-shelled turtles and penguins. During the ascent, turtles actively swam with a stroke frequency of 0.30 Hz but with a low vertical velocity (∼0.40 ms(-1)) and a low pitch angle (∼+26 deg). Turtles might avoid succumbing to decompression sickness ('the bends') by ascending slowly to the surface. In addition, we suggest that the low body temperature of this marine ectotherm compared with that of endotherms might help reduce the risk of bubble formation by increasing the solubility of nitrogen in the blood. This physiological advantage, coupled with several behavioural and physical adaptations, might explain the particular ecological niche the leatherback turtle occupies among marine reptiles.

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Air-breathing animals diving to forage can optimize time underwater by diving with just enough oxygen for the projected performance underwater. By so doing they surface with minimal body oxygen levels, which leads to maximal rates of oxygen uptake. We examined whether imperial cormorants Phalacrocorax atriceps adhere to this by examining dive:pause ratios in birds diving for extended, continuous periods to constant depths, assuming that the oxygen used underwater was exactly replenished by the periods at the surface. Examination of the cumulative time spent in surface pauses relative to the cumulative time spent in diving showed that surface pauses increase according to a power curve function of time spent in the dive or water depth. In a simplistic model we considered the rate at which birds expended energy underwater to be constant and that the rate of oxygen replenishment during the surface pause was directly proportional to the oxygen deficit. We then worked out values for the rate constant for the surface pause before using this constant to examine bird body oxygen levels immediately pre- and post dive. The model predicted that imperial cormorants do not submerge with just enough oxygen to cover their projected dive performance but rather dive with substantial reserves, although these reserves decrease with increasing dive depth/duration. We speculate that these oxygen reserves may be used to enhance bird survival when rare events, such as the appearance of predators or discovery of large prey requiring extended handling time, occur. The form of the oxygen saturation curve over time at the surface means that the time costs for maintaining constant oxygen reserves become particularly onerous for long, deep dives, so the observed decrease in reserves with increasing dive duration is expected in animals benefiting by optimizing for time.

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