Feeding–Danger Trade-Offs Underlie Stopover Site Selection by Migrants

To migrate successfully, birds need to store adequate fat reserves to fuel each leg of the journey. Migrants acquire their fuel reserves at stopover sites; this often entails exposure to predators. Therefore, the safety attributes of sites may be as important as the feeding opportunities. Furthermore, site choice might depend on fuel load, with lean birds more willing to accept danger to obtain good feeding. Here, we evaluate the factors underlying stopover-site usage by migrant Western Sandpipers (Calidris mauri) on a landscape scale. We measured the food and danger attributes of 17 potential stopover sites in the Strait of Georgia and Puget Sound region. We used logistic regression models to test whether food, safety, or both were best able to predict usage of these sites by Western Sandpipers. Eight of the 17 sites were used by sandpipers on migration. Generally, sites that were high in food and safety were used, whereas sites that were low in food and safety were not. However, dangerous sites were used if there was ample food abundance, and sites with low food abundance were used if they were safe. The model including both food and safety best-predicted site usage by sandpipers. Furthermore, lean sandpipers used the most dangerous sites, whereas heavier birds (which do not need to risk feeding in dangerous locations) used safer sites. This study demonstrates that both food and danger attributes are considered by migrant birds when selecting stopover sites, thus both these attributes should be considered to prioritize and manage stopover sites for conservation.

Prioritizing tropical habitats for long-distance migratory songbirds: an assessment of habitat quality at a stopover site in Colombia

Long-distance migratory birds are declining globally and migration has been identified as the primary source of mortality in this group. Despite this, our lack of knowledge of habitat use and quality at stopovers, i.e., sites where the energy for migration is accumulated, remains a barrier to designing appropriate conservation measures, especially in tropical regions. There is therefore an urgent need to assess stopover habitat quality and concurrently identify efficient and cost-effective methods for doing so. Given that fuel deposition rates directly influence stopover duration, departure fuel load, and subsequent speed of migration, they are expected to provide a direct measure of habitat quality and have the advantage of being measurable through body-mass changes. Here, we examined seven potential indicators of quality, including body-mass change, for two ecologically distinct Neotropical migratory landbirds on stopover in shade-coffee plantations and tropical humid premontane forest during spring migration in Colombia: (1) rate of body-mass change; (2) foraging rate; (3) recapture rate; (4) density; (5) flock size; (6) age and sex ratios; and (7) body-mass distribution. We found higher rates of mass change in premontane forest than in shade-coffee in Tennessee Warbler Oreothlypis peregrina, a difference that was mirrored in higher densities and body masses in forest. In Gray-cheeked Thrush Catharus minimus, a lack of recaptures in shade-coffee and higher densities in forest, also suggested that forest provided superior fueling conditions. For a reliable assessment of habitat quality, we therefore recommend using a suite of indicators, taking into account each species’ ecology and methodological considerations. Our results also imply that birds stopping over in lower quality habitats may spend a longer time migrating and require more stopovers, potentially leading to important carryover effects on reproductive fitness. Evaluating habitat quality is therefore imperative prior to defining the conservation value of newly identified stopover regions.

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.

Is re-feeding efficiency compromised by prolonged starvation during aestivation in the green striped burrowing frog, Cyclorana alboguttata?

We examined several morphological parameters of the gastrointestinal tract, digesta passage rates, and nutrient assimilation efficiencies of Green-striped Burrowing frogs (Cyclorana alboguttata) following prolonged fasting during three months of aestivation and compared these with frogs that had been continuously fed. Whole animal digesta passage rates were significantly reduced following three months aestivation as a result of a decreased digesta evacuation rate from the stomach. Furthermore, food was selectively retained in the small intestine for an increased time following three months of aestivation. Overall digestibility of food and nitrogen, carbon, and energy extraction efficiencies were not significantly different from control values following three months of aestivation. These findings suggest that C. alboguttata employs reduced digesta passage rates so as to maximize nutrient assimilation efficiency following prolonged food deprivation during aestivation. (C) 2003 Wiley-Liss, Inc.

Nonbreeding Eastern Curlews Numenius madagascariensis Do Not Increase the Rate of Intake or Digestive Efficiency before Long-Distance Migration because of an Apparent Digestive Constraint

The possibility of premigratory modulation in gastric digestive performance was investigated in a long-distance migrant, the eastern curlew (Numenius madagascariensis), in eastern Australia. The rate of intake in the curlews was limited by the rate of digestion but not by food availability. It was hypothesized that before migration, eastern curlews would meet the increased energy demand by increasing energy consumption. It was predicted that (1) an increase in the rate of intake and the corresponding rate of gastric throughput would occur or (2) the gastric digestive efficiency would increase between the mid-nonbreeding and premigratory periods. Neither crude intake rate (the rate of intake calculated including inactive pauses; 0.22 g DM [grams dry mass] or 3.09 kJ min(-1)) nor the rate of gastric throughput (0.15 g DM or 2.85 kJ min(-1)) changed over time. Gastric digestive efficiency did not improve between the periods (91%) nor did the estimated overall energy assimilation efficiency (63% and 58%, respectively). It was concluded that the crustacean-dominated diet of the birds is processed at its highest rate and efficiency throughout a season. It appears that without a qualitative shift in diet, no increase in intake rate is possible. Accepting these findings at their face value poses the question of how and over what time period the eastern curlews store the nutrients necessary for the ensuing long, northward nonstop flight.

The impact of prolonged fasting during aestivation on the structure of the small intestine in the green-striped burrowing frog, Cyclorana alboguttata

The effects of short-term fasting and prolonged fasting during aestivation on the morphology of the proximal small intestine and associated organs were investigated in the green-striped burrowing frog, Cyclorana alboguttata (Anura: Hylidae). Animals were fasted for 1 week while active or for 3-9 months during aestivation. Short-duration fasting (1 week) had little effect on the morphology of the small intestine, whilst prolonged fasting during aestivation induced marked enteropathy including reductions in intestinal mass, length and diameter, longitudinal fold height and tunica muscularis thickness. Enterocyte morphology was also affected markedly by prolonged fasting: enterocyte cross-sectional area and microvillous height were reduced during aestivation, intercellular spaces were visibly reduced and the prevalence of lymphocytes amongst enterocytes was increased. Mitochondria and nuclei were also affected by 9 months of aestivation with major disruptions to mitochondrial cristae and increased clumping of nuclear material and increased infolding of the nuclear envelope. The present study demonstrates that the intestine of an aestivating frog responds to prolonged food deprivation during aestivation by reducing in size, presumably to reduce the energy expenditure of the organ.

Migration routes and stopover sites of Black-necked Cranes determined by satellite tracking

Because their breeding and wintering areas are in remote locations, little is known about the biology of Black-necked Cranes (Grus nigricollis), including their migratory behavior. Using satellite telemetry, we monitored the migration of Black-necked Cran

Site Safety and Food Affect Movements of Semipalmated Sandpipers (Calidris pusilla) Migrating Through the Upper Bay of Fundy

The upper Bay of Fundy is a critical stopover site for Semipalmated Sandpipers (Calidris pusilla) during their fall migration. However, little is known about factors that influence selection of feeding and roosting sites by these birds, or the extent to which birds move between different sites during their time in the region. Using radio-telemetry, we studied movement patterns, examined habitat use, and tested hypotheses associated with factors influencing foraging and roost-site selection. Movements of radio-tagged sandpipers were tracked in the upper Bay of Fundy in August 2004 and 2005. In 2004, sandpipers from the Minas Basin, Nova Scotia and Chignecto Bay, New Brunswick and Nova Scotia, were tracked, and in 2005, sandpipers were tracked only in Chignecto Bay. Sandpipers were highly mobile in both the Minas Basin 2004 and Chignecto Bay 2005, making daily movements of up to 20 km between foraging and roosting sites, although very little movement was detected in Chignecto Bay in 2004. Migrating sandpipers appeared to select foraging sites based on relative safety, as measured by distance to cover, provided that these sites offered an adequate food supply. Similarly, roosting sandpipers preferred sites that were far from nearby trees that might offer cover to predators. This preference for safe sites became more apparent later in their stay in the Bay of Fundy, when birds were heavier and, therefore, possibly more vulnerable to predation. Semipalmated Sandpipers appear to be flexible during their time in the upper Bay of Fundy, displaying year-to-year and site-to-site variability in movement and mudflat usage. Therefore, multiple, synchronized population counts should be conducted at known roost sites in order to more accurately estimate Semipalmated Sandpiper abundance in this region. Furthermore, in a highly dynamic system where food can be variable, landscape features such as distance to cover may be important factors to consider when selecting candidate sites for shorebird conservation measures.

Flexibility of Continental Navigation and Migration in European Mallards

The ontogeny of continent-wide navigation mechanisms of the individual organism, despite being crucial for the understanding of animal movement and migration, is still poorly understood. Several previous studies, mainly conducted on passerines, indicate that inexperienced, juvenile birds may not generally correct for displacement during fall migration. Waterbirds such as the mallard (Anas platyrhynchos, Linnaeus 1758) are more flexible in their migration behavior than most migratory songbirds, but previous experiments with waterbirds have not yet allowed clear conclusions about their navigation abilities. Here we tested whether immature mallard ducks correct for latitudinal displacement during fall migration within Europe. During two consecutive fall migration periods, we caught immature females on a stopover site in southeast Sweden, and translocated a group of them ca. 1,000 km to southern Germany. We followed the movements of the ducks via satellite GPS-tracking and observed their migration decisions during the fall and consecutive spring migration. The control animals released in Ottenby behaved as expected from banding recoveries: they continued migration during the winter and in spring returned to the population's breeding grounds in the Baltics and Northwest Russia. Contrary to the control animals, the translocated mallards did not continue migration and stayed at Lake Constance. In spring, three types of movement tactics could be observed: 61.5% of the ducks (16 of 26) stayed around Lake Constance, 27% (7 of 26) migrated in a northerly direction towards Sweden and 11.5% of the individuals (3 of 26) headed east for ca. 1,000 km and then north. We suggest that young female mallards flexibly adjust their migration tactics and develop a navigational map that allows them to return to their natal breeding area.

Insights from the eco-physiological book of records : Bewick's swans outperform the canonical intake-maximizing vertebrate

During periods of high energy demand an animal may be constrained by a physiological maximum to its energy intake rate. Predictions by allometric equations describing this maximum for endotherms were significantly surpassed during a few recent laboratory experiments on birds and mammals, being given access to food 24 h day^-1. How relevant this is in the field remains to be assessed. We predicted that Bewick’s swans Cygnus columbianus bewickii might surpass this maximum during stopover on their migration. We determined intake rate by measuring initial and final biomass density, and dividing the biomass difference by the feeding time required to reach this difference. This feeding time was given by the functional response. After conversion to daily energy intake rates, these exceeded the previously assumed maximum on two of the three stopover sites studied. The exception was a stopover site where daily foraging time was limited by the tidal cycle. Our study confirms that intake rates may exceed the formerly generally supposed maximum under natural conditions when foraging is possible day and night.

Intake rates, stochasticity, or onset of spring : what aspects of food availability affect spring migration patterns in Pink-footed Geese Anser brachyrhynchus?

Long-distance bird migration consists of several flight episodes interrupted by a series of resting and refuelling periods on stopover sites. We assessed the role of food availability as the determinant of staging decisions focusing on the following three aspects of food availability: intake rates, stochasticity in intake rates and onset of spring. Using stochastic dynamic modelling, we investigated their impact on staging times and expected fitness. Subsequently, we compared relations in the use of the stopover sites as predicted by the model with empirical data of the Svalbard-breeding population of Pink-footed Goose Anser brachyrhynchus collected in the period 1990–2002. Our results indicate that, for the case of Pink-footed Geese, spring phenology determines a major part of the migration schedule. In contrast to our expectations, intake rates were generally only of minor importance; however, when approaching the breeding grounds their significance increased. Expected fitness at arrival on the breeding grounds showed that the geese can compensate for changes in a broad range of food availability and also cope with varying degrees of stochasticity. However, declining intake rates at the last stopover site or very late onsets of spring clearly decreased fitness. As predicted by the model, the use of stopover sites was interdependent – from empirical data we derived negative relationships between the staging durations of subsequent sites. These results lend credit to an integrated spatially explicit approach focussing on multiple stopover site characteristics when attempting to improve our understanding of bird migration.

Stochastic atmospheric assistance and the use of emergency staging sites by migrants

Numerous animals move vast distances through media with stochastic dynamic properties. Avian migrants must cope with variable wind speeds and directions en route, which potentially jeopardize fine-tuned migration routes and itineraries. We show how unpredictable winds affect flight times and the use of an intermediate staging site by red knots (Calidris canutus canutus) migrating from west Africa to the central north Siberian breeding areas via the German Wadden Sea. A dynamic migration model incorporating wind conditions during flight shows that flight durations between Mauritania and the Wadden Sea vary between 2 and 8 days. The number of birds counted at the only known intermediate staging site on the French Atlantic coast was strongly positively correlated with simulated flight times. In addition, particularly light-weight birds occurred at this location. These independent results support the idea that stochastic wind conditions are the main driver of the use of this intermediate stopover site as an emergency staging area. Because of the ubiquity of stochastically varying media, we expect such emergency habitats to exist in many other migratory systems, both airborne and oceanic. Our model provides a tool to quantify the effect of winds and currents en route.

Ecological factors associated with the breeding and migratory phenology of high-latitude breeding western sandpipers

Environmental conditions influence the breeding and migratory patterns of many avian species and may have particularly dramatic effects on long-distance migrants that breed at northern latitudes. Environment, however, is only one of the ecological variables affecting avian phenology, and recent work shows that migration tactics may be strongly affected by changes in predator populations. We used long-term data from 1978 to 2000 to examine the interactions between snowmelt in western Alaska in relation to the breeding or migration phenologies of small shorebirds and their raptor predators. Although the sandpipers' time of arrival at Alaskan breeding sites corresponded with mean snowmelt, late snowmelts did delay breeding. These delays, however, did not persist to southward migration through British Columbia, likely due to the birds' ability to compensate for variance in the length of the breeding season. Raptor phenology at an early stopover site in British Columbia was strongly related to snowmelt, so that in years of early snowmelt falcons appeared earlier during the sandpipers' southbound migration. These differential effects indicate that earlier snowmelt due to climate change may alter the ecological dynamics of the predator-prey system.

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.