Phil F. Battley

Breeding latitude drives individual schedules in a trans-hemispheric migrant bird

Despite clear benefits of optimal arrival time on breeding grounds, migration schedules may vary with an individual birds innate quality, non-breeding habitat or breeding destination. Here, we show that for the bar-tailed godwit (Limosa lapponica baueri), a shorebird that makes the longest known non-stop migratory flights of any bird, timing of migration for individual birds from a non-breeding site in New Zealand was strongly correlated with their specific breeding latitudes in Alaska, USA, a 16,000-18,000 km journey away. Furthermore, this variation carried over even to the southbound return migration, 6 months later, with birds returning to New Zealand in approximately the same order in which they departed. These tightly scheduled movements on a global scale suggest endogenously controlled routines, with breeding site as the primary driver of temporal variation throughout the annual cycle.

Empirical evidence for differential organ reductions during trans–oceanic bird flight

Since the early 1960s it has been held that migrating birds deposit and use only fat as fuel during migratory flight, with the non–fat portion of the body remaining homeostatic. Recent evidence from field studies has shown large changes in organ sizes in fuelling birds, and theory on fuel use suggests protein may be a necessary fuel during flight. However, an absence of information on the body condition of migrants before and after a long flight has hampered understanding of the dynamics of organs during sustained flight. We studied body condition in a medium–sized shorebird, the great knot (Calidris tenuirostris), before and after a flight of 5400 km from Australia to China during northward migration. Not only did these birds show the expected large reduction in fat content after migration, there was also a decrease in lean tissue mass, with significant decreases in seven organs. The reduction in functional components is reflected in a lowering of the basal metabolic rate by 46%. Recent flight models have tried to separate the ‘flexible’ part of the body from the constant portion. Our results suggest that apart from brains and lungs no organs are homeostatic during long–distance flight. Such organ reductions may be a crucial adaptation for long–distance flight in birds.

Scale and intensity of intertidal habitat use by knots Calidris canutus in the Western Wadden Sea in relation to food, friends and foes

Abstract In August–October 1988–1992 we studied the distribution and abundance of knots Calidris canutus around Griend in the western Wadden Sea, and the extent to which these can be explained by benthic prey availability and presence of avian predators. Numbers in the nonbreeding season showed monthly averages of 10 000 to 25 000 birds. Over 100 000 knots were recorded on three occassions. Knots feed in large flocks, individual birds usually experiencing 4 000 to 15 000 flock-mates. The Siberian-breeding/west-African wintering canutus subspecies passed through in late July and early August. Otherwise the Greenlandic/Canadian breeding islandica subspecies was present. Over the period 1964–1992 there were no clear trends in the number of knots, but canutus -knots were particularly abundant in July–August 1991, whereas in 1992 both subspecies were absent. Macoma balthica was the preferred prey of both subspecies. Hydrobia ulvae, Mytilus edulis and Cerastoderma edule were eaten when Macoma was absent close to the surface of the sediment. As Macoma buried deeper from July onwards, canutus faced better average feeding conditions than islandica later in the year. The spatial distributon of knots feeding on the intertidal flats around Griend was best explained by the harvestable biomass of the prevalent prey species in a particular year and season, i.e. Macoma (main prey when their harvestable biomass densities were greater than ca 0.8 g AFDM per m 2 ) and Cerastoderma , and by the avoidance of situations where they run the risk of attack by bird-eating birds. Flocks of knots covered most of the intertidal flats in the Western Dutch Wadden Sea in a couple of tidal cycles. This is about 800 km 2 , much larger than the equivalent area used by knots on their wintering grounds in Mauritania (10–15 km 2 ), a difference that is correlated with prey spectrum, prey availability and predictability.

Is Long‐Distance Bird Flight Equivalent to a High‐Energy Fast? Body Composition Changes in Freely Migrating and Captive Fasting Great Knots

We studied changes in body composition in great knots, Calidris tenuirostris, before and after a migratory flight of 5,400 km from northwest Australia to eastern China. We also took premigratory birds into captivity and fasted them down to their equivalent arrival mass after migration to compare organ changes and nutrient use in a low‐energy‐turnover fast with a high‐energy‐turnover fast (migratory flight). Migrated birds were as economical as any fasting animal measured yet at conserving protein: their estimated relative protein contribution (RPC) to the energy used was 4.0%. Fasted birds had an estimated RPC of 6.8% and, consequently, a much lower lean mass and higher fat content for an equivalent body mass than migrated birds. Lean tissue was catabolized from most organs in both groups, except the brain. Furthermore, a principal components biplot showed that individuals were grouped primarily on the basis of overall organ fat or lean tissue content rather than by the size of specific organs. This indicates that organ changes during migratory flight are similar to those of a low‐energy fast, although the length of the fast in this study probably accentuated organ reductions in some functional groups. Whether the metabolic characteristics of a flying migratory fast follow the three‐phase model described in many inactive fasting animals is unclear. We have some evidence for skeletal fat being catabolized without phase 3 of a fast having been reached.

Reinterpretation of gizzard sizes of red knots world-wide emphasises overriding importance of prey quality at migratory stopover sites

The size of digestive organs can be rapidly and reversibly adjusted to ecological circumstances, but such phenotypic flexibility comes at a cost. Here, we test how the gizzard mass of a long-distance migrant, the red knot (Calidris canutus), is adjusted to (i) local climate, (ii) prey quality and (iii) migratory fuelling demands. For eight sites around the world (both wintering and stopover sites), we assembled data on gizzard masses of free-living red knots, the quality of their prey and the local climate. Using an energetic cost–benefit approach, we predicted the gizzard size required for fastest fuelling (net rate-maximization, i.e. expected during migration) and the gizzard size required to balance daily energy budgets (satisficing, expected in wintering birds) at each site. The measured gizzards matched the net rate-maximizing predictions at stopover sites and the satisficing predictions at wintering sites. To our surprise, owing to the fact that red knots selected stopover sites with prey of particularly high quality, gizzard sizes at stopovers and at wintering sites were nevertheless similar. To quantify the benefit of minimizing size changes in the gizzard, we constructed a model incorporating the size-dependent energy costs of maintaining and carrying a gizzard. The model showed that by selecting stopovers containing high-quality prey, metabolic rates are kept at a minimum, potentially reducing the spring migratory period by a full week. By inference, red knots appear to time their stopovers so that they hit local peaks in prey quality, which occur during the reproductive seasons of the intertidal benthic invertebrates.

Consistent annual schedules in a migratory shorebird

Many migratory birds start prebreeding moult and premigratory fuelling some months before the breeding season and face severe time constraints, while travelling up to 15 000 km between non-breeding and breeding grounds. Shorebirds typically leave Southern Hemisphere non-breeding areas over a 3–4 week period, but whether they benefit from interannually consistent timing of departure is unknown. Here, I show that individual bar-tailed godwits (Limosa limosa baueri) from New Zealand are highly consistent in their migratory scheduling. Most birds left within the same week each year (between-year repeatability, r, of 0.83) and adult males, which moult into a bright breeding plumage, were also highly repeatable in the extent of their prebreeding moult (r=0.86). This is consistent with the hypothesis that birds have individually optimized migration schedules. Within adult males, but not females, smaller birds tended to migrate earlier than large birds. Whether this reflects differences in size-related migration speed, optimal breeding time at different sites or size-related natural or sexual selection pressures, remains unknown.

High-tide habitat choice: insights from modelling roost selection by shorebirds around a tropical bay

High tides force shorebirds from intertidal feeding areas to sites known as roosts. We investigated the roost selection of great knots, Calidris tenuirostris, and red knots, Calidris canutus, on a tropical coastline in northwestern Australia, assessing several roost attributes and recording the frequency of use of each site through automatic radiotelemetry. To model roost choice we used two approaches: (1) conditional logistic regression models that assumed roost selection to be a trade-off based on a probabilistic assessment of several environmental characteristics; and (2) bounds-based models that assumed that birds selected the nearest roost site to their feeding grounds, provided that threshold values for certain environmental characteristics were met. Bounds-based models were more effective, and we suggest that they offer a closer approach to real roost choice mechanisms. By day, roost choice was affected by distance from the feeding area and microclimate; birds selected nearby roosts where they could stand on cool, wet substrates. Different roost selection criteria were used at night when birds chose safer, but more distant, roosts. Models that assumed that roost choice was influenced by recent experience of roost sites performed better than models that assumed constant assessment of roost quality. This effect was significant only at night, suggesting that memory was used more when information on roost quality was limited. Evidence that roost availability may influence selection of foraging areas is also presented. Our results suggest that shorebirds select roosts by using simple mechanisms, making roost choice models a potentially valuable tool in conservation planning.

Absolute consistency: Individual versus population variation in annual-cycle schedules of a long-distance migrant bird

Flexibility in scheduling varies throughout an organism’s annual cycle, reflecting relative temporal constraints and fitness consequences among life-history stages. Time-selection can act at different scales, either by limiting the range of alternative strategies in the population, or by increasing the precision of individual performance. We tracked individual bar-tailed godwits Limosa lapponica baueri for two full years (including direct observation during non-breeding seasons in New Zealand and geolocator tracking of round-trip migrations to Alaska) to present a full annual-cycle view of molt, breeding, and migration schedules. At both population and individual scales, temporal variation was greater in post-breeding than pre-breeding stages, and greater in molts than in movements, but schedules did not tighten across successive stages of migration toward the breeding grounds. In general, individual godwits were quite consistent in timing of events throughout the year, and repeatability of pre-breeding movements was particularly high (r = 0.82–0.92). However, we demonstrate that r values misrepresent absolute consistency by confounding inter- and intra-individual variation; the biological significance of r values can only be understood when these are considered separately. By doing so, we show that some stages have considerable tolerance for alternative strategies within the population, whereas scheduling of northbound migratory movements was similar for all individuals. How time-selection simultaneously shapes both individual and population variation is central to understanding and predicting adaptive phenological responses to environmental change.

Basal Metabolic Rate Declines During Long-Distance Migratory Flight in Great Knots

Abstract Great Knots (Calidris tenuirostris) make one of the longest migratory flights in the avian world, flying almost 5500 km from Australia to China during northward migration. We measured basal metabolic rate (BMR) and body composition in birds before and after this flight and found that BMR decreased 42%. The mass-specific BMR based on lean mass decreased 33%. We also starved a group of pre-migratory Great Knots in captivity to determine whether they showed the same reduction in BMR without having undergone the hard work of flight. The captive birds showed a similar range and reduction of BMR values as the wild birds. Exponents of relationships between BMR and body mass in different comparisons were high, indicating large changes in BMR as a function of body mass. Analysis of the body composition of ten wild and three captive birds found that the flight muscle mass and intestine mass positively correlated with BMR. La Tasa Metabolica Basal Disminuye durante Vuelos Migratorios de Larga Distancia en Calidris tenuirostris Resumen. Calidris tenuirostris realiza uno de los vuelos migratorios más largos entre las aves, volando desde Australia hasta China durante la migración al norte. Medimos la tasa metabólica basal (TMB) y la composición corporal en aves antes y después del vuelo y encontramos que TMB se reduce en 42%. La TMB específica por masa, o sea la TMB corregida por masa magra, se redujo en 33%. También expusimos un grupo premigratorio de Calidris tenuirostris a un periodo de ayuno en cautiverio, para determinar si mostraban una tasa de reducción similar en la TMB sin haber soportado la dura tarea de volar. Las aves en cautiverio mostraron un rango y una reducción de los valores de la TMB similares a los de las aves en libertad. Los exponentes de la relación entre TMB y masa corporal en diferentes comparaciones fueron altos, indicando grandes cambios de la TMB en función de la masa corporal. El análisis de la composición corporal de diez aves libres y tres en cautiverio mostró que la masa de los músculos del vuelo y la masa de los intestinos esta positivamente correlacionada con la TMB.

Body composition and flight ranges of Bar-tailed Godwits (Limosa lapponica baueri) from New Zealand

Abstract Body composition analysis was performed on 37 Bar-tailed Godwits (Limosa lapponica baueri) that had been illegally shot during migratory fueling in northern New Zealand in March 1992. Adults (n = 35) were heavy (442–721 g), with fat loads of 30–45% of body mass. Two first-year males had only 5.4% and 8.9% fat, respectively, and had 13% less lean (fat-free dry) tissue than the average adult male. Males and females scaled along the same size axes, with little overlap between the sexes (females were larger). Larger birds carried more fat, but fat loads did not differ systematically with size (and, therefore, sex). Fatter birds (after accounting for body size) had larger flight muscle and lung masses, implying premigratory hypertrophy. Males that were not in body molt were fatter, but less colorful, than molting birds. They were also longer-winged and had larger testes but smaller intestines and salt glands, which suggests that they may belong to a subpopulation with a different fueling and molt strategy. Comparison between L. l. baueri and a smaller subspecies (L. l. taymyrensis) indicated that although the two subspecies scaled along the same lean mass-size axes, L. l. baueri deposited much more fat for their body size (1.7 × more than the equivalent-sized L. l. taymyrensis). On the basis of estimated travel costs for L. l. taymyrensis, predicted flight ranges for L. l. baueri ranged from 6,000 to 8,600 km, depending on whether wind assistance is assumed. That suggests that direct flights from New Zealand to southeastern or eastern Asia are possible (e.g. Philippines, 7,200 km; Japan, 8,700; South Korea, 9,100 km). Composition Corporelle et Distance de Vol chez Limosa lapponica baueri de Nouvelle-Zélande