Ontogeny of long distance migration

The movements of some long-distance migrants are driven by innate compass headings that they follow on their first migrations (e.g., some birds and insects), while the movements of other first-time migrants are learned by following more experienced conspecifics (e.g., baleen whales). However, the overall roles of innate, learned, and social behaviors in driving migration goals in many taxa are poorly understood. To look for evidence of whether migration routes are innate or learned for sea turtles, here for 42 sites around the world we compare the migration routes of >400 satellite-tracked adults of multiple species of sea turtle with ∼45 000 Lagrangian hatchling turtle drift scenarios. In so doing, we show that the migration routes of adult turtles are strongly related to hatchling drift patterns, implying that adult migration goals are learned through their past experiences dispersing with ocean currents. The diverse migration destinations of adults consistently reflected the diversity in sites they would have encountered as drifting hatchlings. Our findings reveal how a simple mechanism, juvenile passive drift, can explain the ontogeny of some of the longest migrations in the animal kingdom and ensure that adults find suitable foraging sites.

Use of long-distance migration patterns of an endangered species to inform conservation planning for the world's largest marine protected area

Large marine protected areas (MPAs), each hundreds of thousands of square kilometers, have been set up by governments around the world over the last decade as part of efforts to reduce ocean biodiversity declines, yet their efficacy is hotly debated. The Chagos Archipelago MPA (640,000 km2) (Indian Ocean) lies at the heart of this debate. We conducted the first satellite tracking of a migratory species, the green turtle (Chelonia mydas), within the MPA and assessed the species' use of protected versus unprotected areas. We developed an approach to estimate length of residence within the MPA that may have utility across migratory taxa including tuna and sharks. We recorded the longest ever published migration for an adult cheloniid turtle (3979 km). Seven of 8 tracked individuals migrated to distant foraging grounds, often ≥1000 km outside the MPA. One turtle traveled to foraging grounds within the MPA. Thus, networks of small MPAs, developed synergistically with larger MPAs, may increase the amount of time migrating species spend within protected areas. The MPA will protect turtles during the breeding season and will protect some turtles on their foraging grounds within the MPA and others during the first part of their long-distance postbreeding oceanic migrations. International cooperation will be needed to develop the network of small MPAs needed to supplement the Chagos Archipelago MPA.

Metabolic constraints on long-distance migration in birds

The flight range of migrating birds depends crucially on the amount of fuel stored by the bird prior to migration or taken up en route at stop-over sites. However, an increase in body mass is associated with an increase in energetic costs, counteracting the benefit of fuel stores. Water imbalance, occurring when water loss exceeds metabolic water production, may constitute another less well recognised problem limiting flight range. The main route of water loss during flight is via the lungs; the rate of loss depends on ambient temperature, relative humidity and ventilatory flow and increases with altitude. Metabolite production results in an increased plasma osmolality, also endangering the proper functioning of the organism during flight. Energetic constraints and water-balance problems may interact in determining several aspects of flight behaviour, such as altitude of flight, mode of flight, lap distance and stop-over duration. To circumvent energetic and water-balance problems, a bird could migrate in short hops instead of long leaps if crossing of large ecological barriers can be avoided. However, although necessitating larger fuel stores and being more expensive, migration by long leaps may sometimes be faster than by short hops. Time constraints are also an important factor in explaining why soaring, which conserves energy and water, occurs exclusively in very large species: small birds can soar at low speeds only. Good navigational skills involving accurate orientation and assessment of altitude and air and ground speed assist in avoiding physiological stress during migration.

Why fly the extra mile? Latitudinal trend in migratory fuel deposition rate as driver of trans-equatorial long-distance migration.

Trans-equatorial long-distance migrations of high-latitude breeding animals have been attributed to narrow ecological niche widths. We suggest an alternative hypothesis postulating that trans-equatorial migrations result from a possible increase in the rate at which body stores to fuel migration are deposited with absolute latitude; that is, longer, migrations away from the breeding grounds surpassing the equator may actually enhance fueling rates on the nonbreeding grounds and therewith the chance of a successful, speedy and timely migration back to the breeding grounds. To this end, we first sought to confirm the existence of a latitudinal trend in fuel deposition rate in a global data set of free-living migratory shorebirds and investigated the potential factors causing this trend. We next tested two predictions on how this trend is expected to impact the migratory itineraries on northward migration under the time-minimization hypothesis, using 56 tracks of high-latitude breeding shorebirds migrating along the East Asian-Australasian Flyway. We found a strong positive effect of latitude on fuel deposition rate, which most likely relates to latitudinal variations in primary productivity and available daily foraging time. We next confirmed the resulting predictions that (1) when flying from a stopover site toward the equator, migrants use long jumps that will take them to an equivalent or higher latitude at the opposite hemisphere; and (2) that from here onward, migrants will use small steps, basically fueling only enough to make it to the next suitable staging site. These findings may explain why migrants migrate "the extra mile" across the equator during the nonbreeding season in search of better fueling conditions, ultimately providing secure and fast return migrations to the breeding grounds in the opposite hemisphere.

What drives long-distance movements in the nomadic Grey Teal Anas gracilis in Australia?

In contrast to northern temperate environments, where day length and temperature changes are obvious proximate cues for movement to resource-rich breeding habitats, the cues for movement used by birds in an often resource-poor, stochastic environment are less obvious. We recorded long-distance movements of 23 Grey Teal Anas gracilis using satellite telemetry for up to 879 days and examined the relationship between those movements and environmental factors, such as heavy rainfall and flooding, at the destination site. We identified 32 long-distance [> 150 km) movements that met our criterion for minimally interrupted flight between origin and destination. Thirteen of these flights coincided with rainfall and/or flooding events up to 1050 km from the origin. However, some ducks moved without any clear beneficial conditions at the destination onto small wetlands in regions with little surface water. The data suggest that there are two types of long-distance movement - ranging and directed. These flights occurred over distances up to 1200 km across the arid inland. The rates and distances of movement suggest that long-distance movements of Grey Teal entail high energy costs as in waterfowl elsewhere. We conclude that the proximate controls of directIn contrast to northern temperate environments, where day length and temperature changes are obvious proximate cues for movement to resource-rich breeding habitats, the cues for movement used by birds in an often resource-poor, stochastic environment are less obvious. We recorded long-distance movements of 23 Grey Teal Anas gracilis using satellite telemetry for up to 879 days and examined the relationship between those movements and environmental factors, such as heavy rainfall and flooding, at the destination site. We identified 32 long-distance (> 150 km) movements that met our criterion for minimally interrupted flight between origin and destination. Thirteen of these flights coincided with rainfall and/or flooding events up to 1050 km from the origin. However, some ducks moved without any clear beneficial conditions at the destination onto small wetlands in regions with little surface water. The data suggest that there are two types of long-distance movement – ranging and directed. These flights occurred over distances up to 1200 km across the arid inland. The rates and distances of movement suggest that long-distance movements of Grey Teal entail high energy costs as in waterfowl elsewhere. We conclude that the proximate controls of directed movements need not be very different from those of their temperate counterparts.ed movements need not be very different from those of their temperate counterparts.

Oceanic long-distance navigation : do experienced migrants use the earth's magnetic field?

Albatrosses and sea turtles are known to perform extremely long-distance journeys between disparate feeding areas and breeding sites located on small, isolated, oceanic islands or at specific coastal sites. These oceanic journeys, performed mainly over or through apparently featureless mediums, indicate impressive navigational abilities, and the sensory mechanisms used are still largely unknown. This research used three different approaches to investigate whether bi-coordinate navigation based on magnetic field gradients is likely to explain the navigational performance of wandering albatrosses in the South Atlantic and Indian Oceans and of green turtles breeding on Ascension Island in the South Atlantic Ocean. The possibility that magnetic field parameters can potentially be used in a bi-coordinate magnetic map by wandering albatrosses in their foraging area was investigated by analysing satellite telemetry data published in the literature. The possibilities for using bi-coordinate magnetic navigation varied widely between different areas of the Southern Oceans, indicating that a common mechanism, based on a bi-coordinate geomagnetic map alone, was unlikely for navigation in these areas. In the second approach, satellite telemetry was used to investigate whether Ascension Island green turtles use magnetic information for navigation during migration from their breeding island to foraging areas in Brazilian coastal waters. Disturbing magnets were applied to the heads and carapaces of the turtles, but these appeared to have little effect on their ability to navigate. The only possible effect observed was that some of the turtles with magnets attached were heading for foraging areas slightly south of the control turtles along the Brazilian coast. In the third approach, breeding female green turtles were deliberately displaced in the waters around Ascension Island to investigate which cues these turtles might use to locate and return to the island; the results suggested that cues transported by wind might be involved in the final stages of navigation.

Influence of ocean currents on long-distance movement of leatherback sea turtles in the Southwest Indian Ocean

Leatherback turtles Dermochelys coriacea spend most of their life in oceanic environments, whose physical and biological characteristics are primarily forged by sea current circulation. Water mass movements can mechanically act on swimming turtles, thus determining their routes, and can differentially distribute their planktonic prey. By integrating satellite tracking data with contemporaneous remote-sensing information, we analysed the post-nesting journeys of 9 leatherbacks with respect to oceanographic surface conditions. Tracked turtles showed large variations in migration routes and in final destinations, apparently without heading for specific foraging areas. Their complex tracks spread over wide regions around South Africa. Leatherbacks were greatly influenced by the currents encountered during their movements, with their trajectories displaying curves or revolutions in the presence of (and in accordance with) rotating water masses. An impressive similarity was observed between large parts of the turtle routes and those of surface drifters tracked in the same regions. Finally, leatherbacks remained associated for long periods with specific oceanographic features, which most probably offered them profitable foraging opportunities. These results agree with previous findings in showing a strong influence of oceanic currents and mesoscale features on the movements of South African leatherbacks, and additionally identify the role of current-related features in causing the observed route variability and in determining high-quality foraging hotspots for leatherbacks moving in the ocean.

Long-distance endozoochorous dispersal of submerged macrophyte seeds by migratory waterbirds in northern Europe - a critical review of possibilities and limitations

We review whether migratory Anatidae, i.e., swans, geese and ducks, could be acting as vectors for dispersal of Zostera, Ruppia and Potamogeton propagules by endozoochory (carrying seeds in their guts). We list six prerequisites that must all be fulfilled, if successful dispersal should occur. Several Anatidae species feed on these macrophytes, and undertake rapid long-distance movements, making dispersal possible. We identify four problems, which in combination leads us to conclude that long-distance dispersal events are likely to be rare. (i) Most long-distance movements are out of phase with the reproductive efforts of the plants, and if birds arrive at sites when plants still bear seeds, they are likely to depart well after seed stocks have been depleted. (ii) Seed transport by birds will usually be uni-directional, from north to south on autumn migrations. (iii) Most of the gut contents of migratory birds are likely to have been discarded within 300 km of departure. (iv) In many cases, birds will arrive in habitats seriously different from those they departed, i.e., any seeds carried along will have low chances of surviving in their new site. We suggest that northbound dispersal by endozoochory can only occur during spring if waterbirds feed on seeds that have not been depleted and remained frozen down or buried in sediments, or during moult- or post-moult migrations. Moult migration takes place in summer in phase with the reproductive efforts of the plants. Also epizoochorous dispersal (external attachment) is subject to restrictions i, ii and iv.

Composition of fuel stores and digestive limitations to fuel deposition rate in the long-distance migratory thrush nightingale, Luscinia luscinia

During their autumn migratory phase, thrush nightingales (Luscinia luscinia) previously starved for 2 d were allowed to refuel under three different ambient temperature conditions (-7 degrees, 7 degrees, and 22 degrees C). During the refueling period, as well as during the preceding control and starvation periods, food intake, body mass, and feces production were monitored. In addition, daily energy expenditure was measured during the refueling period. The compilation of the energy balance during the refueling period revealed an energy density of the deposited tissue of 33.6 kJ g-1. Assuming that the deposited tissue consists of fat and protein exclusively, with energy densities of 39.6 and 5.5 kJ g-1 wet mass, respectively, we estimated the deposited tissue to consist of 82% fat and 18% wet protein (6% dry protein and 12% water). Nitrogen balances during control, starvation, and refueling phases and during a period of prolonged and complete starvation indicated that 5% of the nutrient stores consisted of dry protein. Our results support recent findings that nutrient stores for migration often contain protein in addition to fat and consequently are 15%-25% less energy rich than pure fat stores. These proteins might be stored as muscle or other functional tissue and may be required to support the extra mass of the stores and/or reflect an incapacity of the metabolic machinery to catabolize far exclusively. Fuel deposition rate was positively related with ambient temperature, whereas food intake rate was unaffected by temperature. These results indicate that the rate of fuel deposition is limited by a ceiling in food intake rate; when this ceiling is reached, fuel deposition rate is negatively affected by daily energy expenditure rate. To a certain extent, the ceiling in food intake rate varies depending on feeding conditions over the previous days. These variations in food intake capacity probably reflect the building and breakdown of gut tissues and/or gut enzyme systems and might be insensible and not evolutionary adaptive. Significant energetic costs, however, are probably associated with the maintenance of gut tissues. It is therefore feasible that changes in digestive capacity are regulated and are directed at energy economization.

Energetics of fattening and starvation in the long-distance migratory garden warbler, Sylvia borin, during the migratory phase

Garden warblers (Sylvia borin) were subjected to starvation trials during their autumnal migratory phase in order to simulate a period of non-stop migration. Before, during and after this treatment the energy expenditure, activity, food intake and body mass of the subjects were monitored. Assimilation efficiency was constant throughout the experiments. The catabolized (during starvation) and deposited body tissue (during recovery) consisted of 73% fat. Basal metabolic rate was decreased during the starvation period and tended to a gradual increase during the recovery period. The reduced basal metabolic rate can possibly be attributed to a reduced size/function of the digestive system, which is consistent with the sub-maximal food intake immediately after resuming the supply of food to the experimental birds. The observed reductions in basal metabolic rate during starvation and activity during recovery can be viewed as adaptations contributing to a higher economization of energy supplies. The experimental birds were unable to eat large quantities of food directly after a period of starvation leading to a comparatively low, or no increase in body mass. Such a slow mass increase is in agreement with observations of migratory birds on arrival at stop-over sites.

A chain is as strong as its weakest link: assessing the consequences of habitat loss and degradation in a long-distance migratory shorebird

The conservation of migratory species represents a major challenge, as they use multiple sites, all contributing in varying degrees in sustaining high survival and reproductive success. There is particular concern for shorebirds of the East Asian-Australasian Flyway (EAAF), where declining numbers of migratory species have mostly been attributed to habitat loss along the East Asian coast. Using a stochastic dynamic programming migration model, we assessed the effect of habitat degradation scenarios along the EAAF on migration behaviour, survival and reproductive success of a long-distance migrating shorebird, the Ruddy Turnstone (Arenaria interpres). Following manipulation of habitat quality through changes in intake rate, we found that changes on the wintering (major non-breeding) ground in South Australia had the highest negative effect on reproductive success and survival. We also identified Taiwan and the Yellow Sea as sites with high importance for reproductive success. Although habitats along the East Asian coastline are currently most threatened from a range of global change processes, we highlight the importance of conserving high-quality shorebird wintering habitat in Australia. This may be of notable importance to trans-equatorial migratory shorebirds, which often make a long non-stop flight from their wintering grounds in order to skip low-latitude sites that typically provide little food.

Range-wide phylogeography of the Little Penguin (Eudyptula minor): evidence of long-distance dispersal

The Little Penguin (Eudyptula minor), a colonial-nesting seabird that is widespread in New Zealand and southern Australia, has high dispersal potential but exhibits regional variation in morphology, coloration, and breeding phenology. We present a distribution-wide survey of mitochondrial DNA variation in the Little Penguin to document phylogeographic relationships and genetic structuring and to test for concordance with intraspecific taxonomy. Phylogeographic structuring was absent among Australian colonies (27 localities, 94 individuals), but the distribution of haplotypes among colonies was significantly nonrandom (ϕST = 0.110, P < 0.01). The Australian individuals exhibited close phylogenetic relationships with a subset of New Zealand birds (4 localities, 22 individuals), whereas the remaining New Zealand birds (20 localities, 106 individuals) were phylogenetically distinct, with ≥7% sequence divergence, and exhibited greater levels of genetic variation and geographic structuring (ϕST = 0.774, P < 0.05). These patterns are consistent with earlier suggestions of an origin in New Zealand followed by recent colonization of Australia and back-dispersal to New Zealand. Extinction and re-establishment processes may have been important factors in the development of genetic structuring across a range of spatiotemporal scales. The genetic data are consistent with suggestions that a single subspecies exists in Australia, but not with the subspecies distributions within New Zealand that have been suggested on the basis of morphology and coloration.