Osmoregulatory ability predicts geographical range size in marine amniotes.

Species that are distributed over wide geographical ranges are likely to encounter a greater diversity of environmental conditions than do narrowly distributed taxa, and thus we expect a correlation between size of geographical range and breadth of physiological tolerances to abiotic challenges. That correlation could arise either because higher physiological capacity enables range expansion, or because widely distributed taxa experience more intense (but spatially variable) selection on physiological tolerances. The invasion of oceanic habitats by amniotic vertebrates provides an ideal system with which to test the predicted correlation between range size and physiological tolerances, because all three lineages that have secondarily moved into marine habitats (mammals, birds, reptiles) exhibit morphological and physiological adaptations to excrete excess salt. Our analyses of data on 62 species (19 mammals, 18 birds, 24 reptiles) confirm that more-widely distributed taxa encounter habitats with a wider range of salinities, and that they have higher osmoregulatory ability as determined by sodium concentrations in fluids expelled from salt-excreting organs. This result remains highly significant even in models that incorporate additional explanatory variables such as metabolic mode, body size and dietary habits. Physiological data thus may help to predict potential range size and perhaps a species' vulnerability to anthropogenic disturbance.

Temporal dynamics in animal community assembly during post-logging succession in boreal forest.

Species assemblages can result from deterministic processes, such as niche differentiation and interspecific interactions, and from stochastic processes, such as random colonisation and extinction events. Although changes in animal communities following disturbances have been widely examined, few studies have investigated the mechanisms structuring communities during ecological succession. We assessed the impact of logging on small mammal and beetle assemblages in landscapes dominated by old-growth boreal forests. Our objectives were to 1) characterize variations in communities during the first 66 years of post-harvest forest succession, 2) determine if there are non-random patterns of species co-occurrence (i.e., deterministic processes), and if there are, 3) establish whether non-random co-occurrences are best explained by habitat attributes or by interspecific interactions. We captured small mammals and beetles along a gradient of forest succession (5-66 years) and in old-growth forest, and characterized key vegetation attributes. First, we tested whether community compositions in clear-cut stands became similar to those in natural stands after 66 years. We then used null models, which were either unconstrained or constrained by habitat attributes, to address the last two objectives and distinguish effects of vegetation attributes from interspecific interactions on community assembly. We showed that beetle assemblages differed in stands 21-30 years post-harvest compared to old-growth forests. In contrast, harvesting did not influence the composition of small mammal communities. Overall, our results suggest that community assembly during forest succession is driven by both stochastic and deterministic processes, the latter being linked to interspecific interactions more strongly than to vegetation attributes.

Climatic influences on the breeding biology of the agile frog (Rana dalmatina).

Severe population declines of amphibians have been shown to be attributed to climate change. Nevertheless, the various mechanisms through which climate can influence population dynamics of amphibians remain to be assessed, notably to disentangle the relative synergetic or antagonistic influences of temperature and precipitations on specific life history stages. We investigated the impact of rainfall and temperature on the egg-clutch abundance in a population of agile frog (Rana dalmatina) during 29 years (1987-2016) on 14 breeding sites located in Brenne Natural Park, France. Specifically, we examined the influence of environmental conditions occurring during five temporal windows of the year cycle corresponding to specific life history stages. Overall, our results suggest that the year-to-year fluctuations of egg-clutch abundances in Brenne Natural Park were partly dependent on local climatic conditions (rainfall and temperature). Climate seemed to influence breeding frogs during the autumn-winter period preceding reproduction. Spring and summer conditions did not influence reproduction. Additionally, we failed to detect effects of climatic conditions on newly metamorphosed individuals. Other factors such as density dependence and inter-specific interactions with introduced predators are likely to play a significant role in reproduction dynamics of the studied frog populations.

Boldness predicts an individual's position along an exploration-exploitation foraging trade-off.

Individuals do not have complete information about the environment and therefore they face a trade-off between gathering information (exploration) and gathering resources (exploitation). Studies have shown individual differences in components of this trade-off but how stable these strategies are in a population and the intrinsic drivers of these differences is not well understood. Top marine predators are expected to experience a particularly strong trade-off as many species have large foraging ranges and their prey often have a patchy distribution. This environment leads these species to exhibit pronounced exploration and exploitation phases but differences between individuals are poorly resolved. Personality differences are known to be important in foraging behaviour but also in the trade-off between exploration and exploitation. Here we test whether personality predicts an individual exploration-exploitation strategy using wide ranging wandering albatrosses (Diomedea exulans) as a model system. Using GPS tracking data from 276 wandering albatrosses, we extract foraging parameters indicative of exploration (searching) and exploitation (foraging) and show that foraging effort, time in patch and size of patch are strongly correlated, demonstrating these are indicative of an exploration-exploitation (EE) strategy. Furthermore, we show these are consistent within individuals and appear stable in the population, with no reproductive advantage. The searching and foraging behaviour of bolder birds placed them towards the exploration end of the trade-off, whereas shy birds showed greater exploitation. This result provides a mechanism through which individual foraging strategies may emerge. Age and sex affected components of the trade-off, but not the trade-off itself, suggesting these factors may drive behavioural compensation to maintain resource acquisition and this was supported by the evidence that there were no fitness consequence of any EE trait nor the trade-off itself. These results demonstrate a clear trade-off between information gathering and exploitation of prey patches, and reveals for the first time that boldness may drive these differences. This provides a mechanism through which widely reported links between personality and foraging may emerge.

Dispersal and Diving Adjustments of the Green Turtle Chelonia mydas in Response to Dynamic Environmental Conditions during Post-Nesting Migration.

In response to seasonality and spatial segregation of resources, sea turtles undertake long journeys between their nesting sites and foraging grounds. While satellite tracking has made it possible to outline their migration routes, we still have little knowledge of how they select their foraging grounds and adapt their migration to dynamic environmental conditions. Here, we analyzed the trajectories and diving behavior of 19 adult green turtles (Chelonia mydas) during their post-nesting migration from French Guiana and Suriname to their foraging grounds off the coast of Brazil. First Passage Time analysis was used to identify foraging areas located off Ceará state of Brazil, where the associated habitat corresponds to favorable conditions for seagrass growth, i.e. clear and shallow waters. The dispersal and diving patterns of the turtles revealed several behavioral adaptations to the strong hydrodynamic processes induced by both the North Brazil current and the Amazon River plume. All green turtles migrated south-eastward after the nesting season, confirming that they coped with the strong counter North Brazil current by using a tight corridor close to the shore. The time spent within the Amazon plume also altered the location of their feeding habitats as the longer individuals stayed within the plume, the sooner they initiated foraging. The green turtles performed deeper and shorter dives while crossing the mouth of the Amazon, a strategy which would help turtles avoid the most turbulent upper surface layers of the plume. These adjustments reveal the remarkable plasticity of this green turtle population when reducing energy costs induced by migration.

The spatial distribution of Mustelidae in France.

We estimated the spatial distribution of 6 Mustelidae species in France using the data collected by the French national hunting and wildlife agency under the "small carnivorous species logbooks" program. The 1500 national wildlife protection officers working for this agency spend 80% of their working time traveling in the spatial area in which they have authority. During their travels, they occasionally detect dead or living small and medium size carnivorous animals. Between 2002 and 2005, each car operated by this agency was equipped with a logbook in which officers recorded information about the detected animals (species, location, dead or alive, date). Thus, more than 30000 dead or living animals were detected during the study period. Because a large number of detected animals in a region could have been the result of a high sampling pressure there, we modeled the number of detected animals as a function of the sampling effort to allow for unbiased estimation of the species density. For dead animals -- mostly roadkill -- we supposed that the effort in a given region was proportional to the distance traveled by the officers. For living animals, we had no way to measure the sampling effort. We demonstrated that it was possible to use the whole dataset (dead and living animals) to estimate the following: (i) the relative density -- i.e., the density multiplied by an unknown constant -- of each species of interest across the different French agricultural regions, (ii) the sampling effort for living animals for each region, and (iii) the relative detection probability for various species of interest.

Spatial variation in age structure among colonies of a marine snake: the influence of ectothermy.

Several tetrapod lineages that have evolved to exploit marine environments (e.g. seals, seabirds, sea kraits) continue to rely upon land for reproduction and, thus, form dense colonies on suitable islands. In birds and mammals (endotherms), the offspring cannot survive without their parents. Terrestrial colonies contain all age classes. In reptiles (ectotherms), this constraint is relaxed, because offspring are independent from birth. Hence, each age class has the potential to select sites with characteristics that favour them. Our studies of sea snakes (sea kraits) in the lagoon of New Caledonia reveal marked spatial heterogeneity in age structure among colonies. Sea krait colonies exhibit the endothermic 'seal-seabird' pattern (mixed-age classes within populations) only where the lagoon is narrow. Where the lagoon is wide, most snake colonies are comprised primarily of a single age cohort. Nurseries are located near the coast, adult colonies offshore and mixed colonies in-between. We suggest that ectothermy allows individuals to utilize habitats that are best suited to their own ecological requirements, a flexibility not available to endothermic marine taxa with obligate parental care.

Is pre-breeding prospecting behaviour affected by snow cover in the irruptive snowy owl? A test using state-space modelling and environmental data annotated via Movebank.

BACKGROUND: Tracking individual animals using satellite telemetry has improved our understanding of animal movements considerably. Nonetheless, thorough statistical treatment of Argos datasets is often jeopardized by their coarse temporal resolution. State-space modelling can circumvent some of the inherent limitations of Argos datasets, such as the limited temporal resolution of locations and the lack of information pertaining to the behavioural state of the tracked individuals at each location. We coupled state-space modelling with environmental characterisation of modelled locations on a 3-year Argos dataset of 9 breeding snowy owls to assess whether searching behaviour for breeding sites was affected by snow cover and depth in an arctic predator that shows a lack of breeding site fidelity. RESULTS: The state-space modelling approach allowed the discrimination of two behavioural states (searching and moving) during pre-breeding movements. Tracked snowy owls constantly switched from moving to searching behaviour during pre-breeding movements from mid-March to early June. Searching events were more likely where snow cover and depth was low. This suggests that snowy owls adapt their searching effort to environmental conditions encountered along their path. CONCLUSIONS: This modelling technique increases our understanding of movement ecology and behavioural decisions of individual animals both locally and globally according to environmental variables.

A space oddity: geographic and specific modulation of migration in Eudyptes penguins.

Post-breeding migration in land-based marine animals is thought to offset seasonal deterioration in foraging or other important environmental conditions at the breeding site. However the inter-breeding distribution of such animals may reflect not only their optimal habitat, but more subtle influences on an individual's migration path, including such factors as the intrinsic influence of each locality's paleoenvironment, thereby influencing animals' wintering distribution. In this study we investigated the influence of the regional marine environment on the migration patterns of a poorly known, but important seabird group. We studied the inter-breeding migration patterns in three species of Eudyptes penguins (E. chrysolophus, E. filholi and E. moseleyi), the main marine prey consumers amongst the World's seabirds. Using ultra-miniaturized logging devices (light-based geolocators) and satellite tags, we tracked 87 migrating individuals originating from 4 sites in the southern Indian Ocean (Marion, Crozet, Kerguelen and Amsterdam Islands) and modelled their wintering habitat using the MADIFA niche modelling technique. For each site, sympatric species followed a similar compass bearing during migration with consistent species-specific latitudinal shifts. Within each species, individuals breeding on different islands showed contrasting migration patterns but similar winter habitat preferences driven by sea-surface temperatures. Our results show that inter-breeding migration patterns in sibling penguin species depend primarily on the site of origin and secondly on the species. Such site-specific migration bearings, together with similar wintering habitat used by parapatrics, support the hypothesis that migration behaviour is affected by the intrinsic characteristics of each site. The paleo-oceanographic conditions (primarily, sea-surface temperatures) when the populations first colonized each of these sites may have been an important determinant of subsequent migration patterns. Based on previous chronological schemes of taxonomic radiation and geographical expansion of the genus Eudyptes, we propose a simple scenario to depict the chronological onset of contrasting migration patterns within this penguin group.

Looking like mother makes mallard ducklings dominant over their siblings.

Colour variation in time and space among animals may affect social relationships such as pairing and dominance interactions. For instance, some birds are naturally sensitive to leg colour, with some colours being more visible or attractive than others. The colour of the leg-rings used to mark birds may thus be related to behavioural and reproductive variables. Most studies have investigated this effect for adults during reproduction, but leg-ring colour may also affect the behaviour of young birds. We tested the potential effect of leg-ring colours on the within-brood dominance hierarchy of mallard (Anas platyrhynchos) ducklings while each brood formed a stable and exclusive family unit with its mother. Ducklings did not acquire a within-brood dominance rank according to the colour of their own ring. This result suggests that mallards may not have a sensory bias for a given colouration. However, ducklings wearing a ring of the same colour as one of the two rings of their mother were dominant over their siblings. We discuss the potential behavioural and methodological implications of this result.

Relationship between reversed sexual dimorphism, breeding investment and foraging ecology in a pelagic seabird, the masked booby.

Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearin-g-the most energy-demanding period--and may be related to evolution of RSD.

Assessment of scale-dependent foraging behaviour in southern elephant seals incorporating the vertical dimension: a development of the First Passage Time method.

1. Identifying the spatial scales at which top marine predators forage is important for understanding oceanic ecosystems. Several methods quantify how individuals concentrate their search effort along a given path. Among these, First-Passage Time (FPT) analysis is particularly useful to identify transitions in movement patterns (e.g. between searching and feeding). This method has mainly been applied to terrestrial animals or flying seabirds that have little or no vertical component to their foraging, so we examined the differences between classic FPT and a modification of this approach using the time spent at the bottom of a dive for characterizing the foraging activity of a diving predator: the southern elephant seal. 2. Satellite relayed data loggers were deployed on 20 individuals during three successive summers at the Kerguelen Islands, providing a total of 72 978 dives from eight juvenile males and nine adult females. 3. Spatial scales identified using the time spent at the bottom of a dive ( = 68.2 +/- 42.1 km) were smaller than those obtained by the classic FPT analysis ( = 104.7 +/- 67.3 km). Moreover, foraging areas identified using the new approach clearly overlapped areas where individuals increased their body condition, indicating that it accurately reflected the foraging activity of the seals. 4. These results suggest that incorporating the vertical dimension into FPT provides a different result to the surface path alone. Close to the Antarctic continent, within the pack-ice, sinuosity of the path could be explained by a high sea-ice concentration (restricting elephant seal movements), and was not necessarily related to foraging activity. 5. Our approach distinguished between actual foraging activity and changes in behaviour induced by the physical environment like sea ice, and could be applied to other diving predators. Inclusion of diving parameters appears to be essential to identify the spatial scale of foraging areas of diving animals.

Does prey capture induce area-restricted search? A fine-scale study using GPS in a marine predator, the wandering albatross.

In a patchy environment, predators are expected to increase turning rate and start an area-restricted search (ARS) when prey have been encountered, but few empirical data exist for large predators. By using GPS loggers with devices measuring prey capture, we studied how a marine predator adjusts foraging movements at various scales in relation to prey capture. Wandering albatrosses use two tactics, sit and wait and foraging in flight, the former tactic being three times less efficient than the latter. During flight foraging, birds caught large isolated prey and used ARS at scales varying from 5 to 90 km, with large-scale ARS being used only by young animals. Birds did not show strong responses to prey capture at a large scale, few ARS events occurred after prey capture, and birds did not have high rates of prey capture in ARS. Only at small scales did birds increase sinuosity after prey captures for a limited time period, and this occurred only after they had caught a large prey item within an ARS zone. When this species searches over a large scale, the most effective search rule was to follow a nearly straight path. ARS may be used to restrict search to a particular environment where prey capture is more predictable and profitable.

At-sea distribution and scale-dependent foraging behaviour of petrels and albatrosses: a comparative study.

1. In order to study and predict population distribution, it is crucial to identify and understand factors affecting individual movement decisions at different scales. Movements of foraging animals should be adjusted to the hierarchical spatial distribution of resources in the environment and this scale-dependent response to environmental heterogeneity should differ according to the forager's characteristics and exploited habitats. 2. Using First-Passage Time analysis, we studied scales of search effort and habitat used by individuals of seven sympatric Indian Ocean Procellariiform species fitted with satellite transmitters. We characterized their search effort distribution and examined whether species differ in scale-dependent adjustments of their movements according to the marine environment exploited. 3. All species and almost all individuals (91% of 122 individuals) exhibited an Area-Restricted Search (ARS) during foraging. At a regional scale (1000s km), foraging ranges showed a large spatial overlap between species. At a smaller scale (100s km, at which an increase in search effort occurred), a segregation in environmental characteristics of ARS zones (where search effort is high) was found between species. 4. Spatial scales at which individuals increased their search effort differed between species and also between exploited habitats, indicating a similar movement adjustment for predators foraging in the same habitat. ARS zones of the two populations of wandering albatross Diomedea exulans (Crozet and Kerguelen) were similar in their adjustments (i.e. same ARS scale) as well as in their environmental characteristics. These two populations showed a weak spatial overlap in their foraging distribution, with males foraging in more southerly waters than females in both populations. 5. This study demonstrates that predators of several species adjust their foraging behaviour to the heterogeneous environment and these scale-dependent movement adjustments depend on both forager and environment characteristics.