Prolonged and flexible primary moult overlaps extensively with breeding in beach-nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6-month breeding season. Moult timing was estimated using the Underhill-Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success. © 2014 British Ornithologists' Union.

Geolocator studies on Ruddy Turnstones Arenaria interpres and Greater Sandplovers Charadrius leschenaultii in the East Asian-Australasia Flyway reveal widely different migration strategies

In 2010, following successful trials with geolocators on Ruddy Turnstones in 2009, a total of 105 units, of four different models, were deployed at five locations on Ruddy Turnstones and Greater Sandplovers. Geolocator retrieval rates were 44% on Ruddy Turnstone and 27% on Greater Sandplover. Complete (59%) and partial (15%) technical failure rates on geolocators were high and were mostly the result of wear and saltwater corrosion. All 30 units from the Swiss Ornithological Institute failed. Only half of the Mk10 and Mk12 units from the British Antarctic Survey produced full migration histories. The northward migration of Ruddy Turnstones was on a narrow path with many birds completing an initial non-stop flight of 7,600 km to Taiwan. Later, most made a stopover in the Yellow Sea. Median migration duration was 39.5 days and average migration speed of the first major leg of the journey (assuming the birds followed the great circle route between stopovers) was 63.4 kph. Southward migration paths showed a much wider spread, ranging from Mongolia to the central Pacific. The latter involved the same bird that had been tracked along this route the previous year. It has now been logged on similar 27,000 km round trips in two successive years. The median duration of southward migration (78 days) was nearly twice that of northward migration and data on average migration speed for just two migration legs indicated that it might be lower, 30 and 40 kph being the values recorded. Greater Sandplovers were only tracked on northward migration but seemed to follow a similar migration strategy with a large initial non-stop flight followed by shorter flights and more regular stopovers. Plans are outlined for further analyses and future deployments of geolocators.

The significance of northern-central Bass Strait in south-eastern Australia as habitat for burrowing seabirds

The present study provides the first complete estimate of the abundance and distribution of burrowing seabirds in northern-central Bass Strait, a key region for breeding seabirds in south-eastern Australia. The estimated total number of breeding burrows in the region in 2008-11 were 755300±32400 (s.e.) burrows of Short-tailed Shearwaters (Ardenna tenuirostris), 26700±3500 of Little Penguins (Eudyptula minor), 19100±2200 of Common Diving-Petrels (Pelecanoides urinatrix) and 4200±2700 of Fairy Prions (Pachyptila turtur). These represent substantial proportions of the total estimated Australian breeding populations of these species: 6% of the total population of Short-tailed Shearwaters, 14% of Little Penguins, 0.4% of Fairy Prions and 13% of Common Diving-Petrels. Based on the number of active burrows, the number of breeding Short-tailed Shearwaters in the region is estimated to have decreased 35% between 1978-80 and 2008-11, equivalent to a decrease of 1.4% per annum between 1980 and 2011. The regional population of Little Penguins, however, appears to have increased substantially over the same period. Identification of population trends of the other species is limited by a lack of previous data. The importance of this area for burrowing seabirds and the substantial decline in numbers of Short-tailed Shearwaters warrants more frequent monitoring of the abundance of seabirds in the region to allow a robust comparison of changes in populations over time as well as the identification of possible causative factors. © BirdLife Australia 2014.

The exception to the rule : Retreating ice front makes Bewick's swans Cygnus columbianus bewickii migrate slower in spring than in autumn

In the vast majority of migratory bird species studied so far, spring migration has been found to proceed faster than autumn migration. In spring, selection pressures for rapid migration are purportedly higher, and migratory conditions such as food supply, daylength, and/or wind support may be better than in autumn. In swans, however, spring migration appears to be slower than autumn migration. Based on a comparison of tundra swan Cygnus columbianus tracking data with long-term temperature data from wheather stations, it has previously been suggested that this was due to a capital breeding strategy (gathering resources for breeding during spring migration) and/or to ice cover constraining spring but not autumn migration. Here we directly test the hypothesis that Bewick's swans Cygnus columbianus bewickii follow the ice front in spring, but not in autumn, by comparing three years of GPS tracking data from individual swans with concurrent ice cover data at five important migratory stop-over sites. In general, ice constrained the swans in the middle part of spring migration, but not in the first (no ice cover was present in the first part) nor in the last part. In autumn, the swans migrated far ahead of ice formation, possibly in order to prevent being trapped by an early onset of winter. We conclude that spring migration in swans is slower than autumn migration because spring migration speed is constrained by ice cover. This restriction to spring migration speed may be more common in northerly migrating birds that rely on freshwater resources. © 2013 The Authors.

New insights from geolocators deployed on waders in Australia

Geolocators were deployed on waders in Australia for a third successive year, in Feb/Apr 2011 including on Eastern Curlew and Sanderling for the first time. Retrieval rates, in the 2011/12 austral summer, varied markedly between species. Technical performance of the geolocators was better than in previous years. However units on Greater Sand Plovers, migrating to breeding grounds in the Gobi Desert, China/Mongolia, again behaved erratically, and exhibited symptoms suggesting extraneous electromagnetic interference. Generally, for each species studied, the results confirm earlier indications that the first step of northward migration from Australia is a long non-stop flight. Subsequent movements to breeding areas are usually shorter with up to three stopovers in SE Asia or Siberia. Similarly southward migration strategies include at least one long nonstop flight, though this is usually the second (or later) leg of the journey. The timing of migration appears to be particularly related to breeding latitude. Eastern Curlews, which breed at relatively southern latitudes, depart from SE Australia from early March, reach the breeding grounds and lay eggs in April, set off on return migration in early June and, after a long stopover in the Yellow Sea, arrive back in SE Australia in early August. In contrast arctic-breeding Ruddy Turnstones do not depart from SE Australia until mid/late April and do not arrive back at their non-breeding locations until October, with the last individuals (which have taken a trans-Pacific route) not returning until late November/early December. Recorded migration speeds (assuming the birds take a great circle route) were quite variable, ranging from 32 to 84 km/h, presumably due to wind conditions. They generally averaged nearer to 50 km/h rather than the 60–70 km/h which waders are known to be capable of achieving and which has been the basis of some past flight range calculations.

Flying, fasting, and feeding in birds during migration: a nutritional and physiological ecology perspective

Unlike exercising mammals, migratory birds fuel very high intensity exercise (e.g., flight) with fatty acids delivered from the adipose tissue to the working muscles by the circulatory system. Given the primary importance of fatty acids for fueling intense exercise, we discuss the likely limiting steps in lipid transport and oxidation for exercising birds and the ecological factors that affect the quality and quantity of fat stored in wild birds. Most stored lipids in migratory birds are comprised of three fatty acids (16:0, 18:1 and 18:2) even though migratory birds have diverse food habits. Diet selection and selective metabolism of lipids play important roles in determining the fatty acid composition of birds which, in turn, affects energetic performance during intense exercise. As such, migratory birds offer an intriguing model for studying the implications of lipid metabolism and obesity on exercise performance. We conclude with a discussion of the energetic costs of migratory flight and stopover in birds, and its implications for bird migration strategies.

Fuelling rates of garganey (Anas querquedula) staging in the Camargue, Southern France, during spring migration

Most species of long-distance migratory birds put on energy stores to fuel their travels. However, recent studies have highlighted the potential costs associated with carrying too much fuel, either through increased predation risk or decreased flight efficiency. Consequently, it is now widely accepted that migratory birds should carry optimal rather than maximum fuel loads. Information from 372 garganey (Anas querquedula) ringed and recaptured at least once during the same spring in the Camargue, southern France, was used to document fuelling rates of individual ducks in relation to environmental variation and individual variation in condition. On average, garganey added very little fuel stores in the Camargue (mean gain per day = 0.33 g, less than 0.5% of mean body-mass in total over an average stay of 5 days). Fuelling rates were negatively correlated with body mass at capture, but it cannot be excluded that this pattern was a statistical artefact. Given their body-mass at ringing, garganey could potentially still fly long distances when they stop in the Camargue. It is therefore likely that the aim of their stay in southern France is more for resting than refuelling, a finding that may have implications for the proper management of stop-over sites.

Migrating swans profit from favourable changes in wind conditions at low altitude

Because energy reserves limit flight range, wind assistance may be of crucial importance for migratory birds. We tracked eight Bewick's swans Cygnus columbianus bewickii, using 95-g satellite transmitters with altimeters and activity sensors, during their spring migration from Denmark to northern Russia in 1996. During the 82 occasions where a swan's location was recorded in flight, average flight altitude was 165 m a.s.1. with a maximum of 759 m a.s.1., despite winds often being more favourable at higher altitudes. We also counted Bewick's swans departing from the Gulf of Finland and subsequently passing an observatory in the next major stop-over area 800 km further north in the White Sea, northern Russia, during the springs of 1994, 1995 and 1996. A comparison of these counts with wind data provided evidence for Bewick's swans using favourable changes in wind conditions to embark on migration. Changes in the numbers of birds arriving in the White Sea correlated best with favourable changes in winds in the Gulf of Finland 1 day earlier. Again, migratory volume showed a correlation with winds at low altitudes only, despite wind conditions for the swans being more favourable at high altitudes. We conclude that the relatively large Bewick's swan tends to gear its migration to wind conditions at low altitude only. We argue that Bewick's swans do not climb to high altitudes because of mechanical and physiological limitations with respect to the generation of power for flight and to avoid rapid dehydration.

Fuel stores of juvenile waders on autumn migration in high arctic Canada

Little is known about the fuel stores that arctic-breeding waders put on before departure from the breeding grounds. During a ship-based expedition to arctic Canada, we caught waders at seven, mainly coastal sites, with-in 68°-76°N and 139°-67°W, from 28 July to 31 August 1999. More than two hundred waders of twelve species were trapped, mainly White-rumped Calidris fuscicollis, Semipalmated C. pusilla, Baird's C. bairdii and Buff-breasted Sandpipers Tryngites subruficollis. The vast majority of the birds were juveniles. Body masses and visual fat stores were low, close to the lowest values found anywhere during the non-breeding season for the different species. The relatively fattest birds were Buff-breasted Sandpipers, but they were still far from their maximum body mass on spring migration. We conclude that juvenile arctic waders depart from their natal areas with only small fuel stores, which is in concordance with a time-minimising migration strategy.

Skipping swans: fuelling rates and wind conditions determine differential use of migratory stopover sites of Bewick's Swans Cygnus bewickii

Some migratory birds refuel at stopover sites that they by-pass on the return trip. In theory, this skipping behaviour is only expected in time-selected migrants when the overflown site is of a lower quality than the departure site. We provide empirical evidence that quality differences in stopover sites are the cause for skipping in Bewick's Swans Cygnus bewickii tracked by satellite telemetry. Two and five complete tracks were recorded in spring and autunm, respectively, showing that the White Sea was visited for c. 2 weeks in spring, but by-passed (or visited for a few days at the most) in autumn. Skipping of the White Sea in autumn was predicted by a dynamic programming model which was based on calculated gain rates during stopover in the Pechora Delta and the White Sea. This prediction was not sensitive to plausible variations in gain rates. Relative to the Pechora Delta the White Sea is a poor site because a large tidal amplitude precludes foraging on the beds of the submerged macrophyte Fennel Pondweed Potamogeton pectinatus during high tide. The dynamic programming model predicted a fast autunm migration. However, the phenology of autunm arrival dates of Bewick's Swans on the wintering grounds revealed that only in three out of ten years a significant number of birds was able to reach the wintering grounds without refuelling. In the other years, unfavourable wind conditions along the Russian/Baltic part of the route prevented such non-stop migration.

Flight costs and fuel composition of a bird migrating in a wind tunnel

We studied the energy and protein balance of a Thrush Nightingale Luscinia luscinia, a small long-distance migrant, during repeated 12-hr long flights in a wind tunnel and during subsequent two-day fueling periods. From the energy budgets we estimated the power requirements for migratory flight in this 26 g bird at 1.91 Watts. This is low compared to flight cost estimates in birds of similar mass and with similar wing shape. This suggests that power requirements for migratory flight are lower than the power requirements for nonmigratory flight. From excreta production during flight, and nitrogen and energy balance during subsequent fueling, the dry protein proportion of stores was estimated to be around 10%. A net catabolism of protein during migratory flight along with that of fat may reflect a physiologically inevitable process, a means of providing extra water to counteract dehydration, a production of uric acid for anti-oxidative purposes, and adaptive changes in the size of flight muscles and digestive organs in the exercising animal.

Flight altitude of trans-Sahara migrants in autumn: a comparison of radar observations with predictions from meteorological conditions and water and energy balance models

Radar observations on the altitude of bird migration and altitudinal profiles of meteorological conditions over the Sahara desert are presented for the autumn migratory period. Migratory birds fly at an average altitude of 1016 m (a.s.l.) during the day and 571 m during the night. Weather data served to calculate flight range using two models: an energy model (EM) and an energy-and-water model (EWM). The EM assumes that fuel supply limits flight range whereas the EWM assumes that both fuel and water may limit flight range. Flight ranges estimated with the EM were generally longer than those with the EWM. This indicates that trans-Sahara migrants might have more problems balancing their water than their energy budget. However, if we assume fuel stores to consist of 70% instead of 100% fat (the remainder consisting of 9% protein and 21% water), predicted flight ranges of the EM and EWM largely overlap. Increased oxygen extraction, reduced flight costs, reduced exhaled air temperature, reduced cutaneous water loss and increased tolerance to water loss are potential physiological adaptations that would improve the water budget in migrants. Both the EM and EWM predict optimal flight altitudes in agreement with radar observations in autumn. Optimal flight altitudes are differently predicted by the EM and EWM for nocturnal spring migration. During spring, the EWM predicts moderately higher and the EM substantially higher flight altitudes than during autumn. EWM predictions are therefore in better agreement with radar observations on flight altitude of migrants over the Negev desert in spring than EM predictions.

Predicting migratory flight altitudes by physiological migration models

Using the altitudinal profiles of wind, temperature, pressure, and humidity in three flight models, we tried to explain the altitudinal distributions of nocturnal migrants recorded by radar above a desert in southern Israel. In the simplest model, only the tailwind component was used as a predictor of the most preferred flight altitude (T model). The energy model (E model) predicted flight ranges according to mechanical power consumption in flapping flight depending on air density and wind conditions, assuming optimal adjustment of airspeed and compensation of crosswinds, and including the influence of mass loss during flight. The energy-water model (EW model) used the same assumptions and parameters as the E model but also included restrictions caused by dehydration. Because wind was by far the most important factor governing altitudinal distribution of nocturnal migrants, differences in predictions of the three models were small. In a first approach, the EW model performed slightly better than the E model, and both performed slightly better than the T model. Differences were most pronounced in spring, when migrants should fly high according to wind conditions, but when climbing and descending they must cross lower altitudes where conditions are better with respect to dehydration. A simplified energy model (Es model) that omits the effect of air density on flight costs explained the same amount of variance in flight altitude as the more complicated E and EW models. By omitting the effect of air density, the Es model predicted lower flight altitudes and thus compensated for factors that generally bias height distributions downward but are not considered in the models (i.e. climb and descent through lower air layers, cost of ascent, and decrease of oxygen partial pressure with altitude). Our results confirm that wind profiles, and thus energy rather than water limitations, govern the altitudinal distribution of nocturnal migrants, even under the extreme humidity and temperature conditions in the trade wind zone.