The development of activity ranges in juvenile Brown Boobies Sula leucogaster

The post‐fledging dependence period is extremely important because it allows young birds the opportunity to develop behavioural skills required for later life. We raised 12 hatchling Brown Boobies Sula leucogaster and attached a miniaturized GPS logger to each bird to examine how flight improves after fledging. The Boobies made daily trips and increased the maximum distance, total distance travelled each day, trip duration and flight speed. Young Boobies seemed to gradually acquire flight skills towards independence.     

The three-dimensional flight of red-footed boobies: adaptations to foraging in a tropical environment?

In seabirds a broad variety of morphologies, flight styles and feeding methods exist as an adaptation to optimal foraging in contrasted marine environments for a wide variety of prey types. Because of the low productivity of tropical waters it is expected that specific flight and foraging techniques have been selected there, but very few data are available. By using five different types of high-precision miniaturized logger (global positioning systems, accelerometers, time depth recorders, activity recorders, altimeters) we studied the way a seabird is foraging over tropical waters. Red-footed boobies are foraging in the day, never foraging at night, probably as a result of predation risks. They make extensive use of wind conditions, flying preferentially with crosswinds at median speed of 38 km h(-1), reaching highest speeds with tail winds. They spent 66% of the foraging trip in flight, using a flap-glide flight, and gliding 68% of the flight. Travelling at low costs was regularly interrupted by extremely active foraging periods where birds are very frequently touching water for landing, plunge diving or surface diving (30 landings h(-1)). Dives were shallow (maximum 2.4 m) but frequent (4.5 dives h(-1)), most being plunge dives. While chasing for very mobile prey like flying fishes, boobies have adopted a very active and specific hunting behaviour, but the use of wind allows them to reduce travelling cost by their extensive use of gliding. During the foraging and travelling phases birds climb regularly to altitudes of 20-50 m to spot prey or congeners. During the final phase of the flight, they climb to high altitudes, up to 500 m, probably to avoid attacks by frigatebirds along the coasts. This study demonstrates the use by boobies of a series of very specific flight and activity patterns that have probably been selected as adaptations to the conditions of tropical waters.     

Social interactions of juvenile brown boobies at sea as observed with animal-borne video cameras

While social interactions play a crucial role on the development of young individuals, those of highly mobile juvenile birds in inaccessible environments are difficult to observe. In this study, we deployed miniaturised video recorders on juvenile brown boobies Sula leucogaster, which had been hand-fed beginning a few days after hatching, to examine how social interactions between tagged juveniles and other birds affected their flight and foraging behaviour. Juveniles flew longer with congeners, especially with adult birds, than solitarily. In addition, approximately 40% of foraging occurred close to aggregations of congeners and other species. Young seabirds voluntarily followed other birds, which may directly enhance their foraging success and improve foraging and flying skills during their developmental stage, or both.     

Sex differences in Little Auk Alle alle parental care: transition from biparental to paternal-only care

Understanding differences in male and female care in biparental care systems can help interpret the selective pressures that shape parental strategies. We examined Little Auk Alle alle parental care at a breeding colony during the chick-rearing and fledging periods by conducting observations on marked, known-sex pairs, and by examining the sex ratio of birds carrying food to the colony. Little Auks transitioned from biparental to mostly paternal-only care during late chick-rearing. Males delivered more meals and spent more time at the colony than females during late chick-rearing. Very few females were present at the colony by the end of chick-rearing and through the fledging period, and all marked parents observed accompanying their chick to sea were male. Chick mass loss prior to fledging was associated with the lack of provisioning by the female parent, rather than a reduction in feeding frequency by both parents. The occurrence of paternal-only care during and after fledging is discussed in relation to physiological, ecological and phylogenetic constraints.     

The post‐fledging period in a tropical bird: patterns of parental care and survival

How environmental conditions affect the timing and extent of parental care is a fundamental question in comparative studies of life histories. The post‐fledging period is deemed critical for offspring fitness, yet few studies have examined this period, particularly in tropical birds. Tropical birds are predicted to have extended parental care during the post‐fledging period and this period may be key to understanding geographic variation in avian reproductive strategies. We studied a neotropical passerine, the western slaty‐antshrike Thamnophilus atrinucha, and predicted greater care and higher survival during the post‐fledging period compared to earlier stages. Furthermore, we predicted that duration of post‐fledging parental care and survival would be at the upper end of the distribution for Northern Hemisphere passerines. Correspondingly, we observed that provisioning continued for 6–12 weeks after fledging. In addition, provisioning rate was greater after fledging and offspring survival from fledging to independence was 75%, greater than all estimates from north‐temperate passerines. Intervals between nesting attempts were longer when the first brood produced successful fledglings compared to nests where offspring died either in the nest or upon fledging. Parents delayed initiating second nests after the first successful brood until fledglings were near independence. Our results indicate that parents provide greater care after fledging and this extended care likely increased offspring survival. Moreover, our findings of extended post‐fledging parental care and higher post‐fledging survival compared to Northern Hemisphere species have implications for understanding latitudinal variation in reproductive effort and parental investment strategies.     

A developmental constraint on the fledging time of birds

We examined the hypothesis that the rate of bone growth limits the minimum fledging time of birds. Previous observations in California gulls indicate that linear growth of wing bones may be the rate limiting factor in wing development. If bone growth is rate limiting, then birds with relatively long bones for their size could be expected to have longer fledging periods than birds with relatively short bones. We tested this by comparing the length of wing bones, relative to body mass, to the relative length of fledging periods among 25 families. The results support the hypothesis. A strong correlation exists between relative fledging period and relative bone length. Species which have relatively long bones for their body size tend to take longer to fly. In contrast, parameters that influence flight style and performance, such as size of the pectoralis muscle and wing loading, show little or no correlation with fledging time. The analysis also indicates that, when altricial and precocial species are considered together, bone length is more highly correlated with fledging time than is body mass or rate of increase in body mass during growth. These observations suggest that linear growth of bones does limit the growth of avian wings and that it is one of the factors that influences the fledging time of birds.     

Nest attendance and foraging movements of northern fulmars rearing chicks at Bjørnøya Barents Sea

We studied several aspects of the foraging ecology of fulmars rearing young chicks on Bjørnøya. To determine precisely the duration of foraging trips during the brooding period, we used an automated logging system that recorded the presence of fulmars fitted with transponders. We also tracked, with satellite transmitters, four parent fulmars during the brooding period, and two after the chick had been left alone. When brooding the chick, fulmars appeared to alternate very rapidly on the nest, with foraging trips lasting on average 8?h. This period appeared constraining for the birds since parents lost mass. The growth of chicks was dependent on the ability of the female (and not the male) to do short foraging trips. At this time birds are foraging at an average distance of 60?km from the colony, with birds concentrating on the shelf around Bjørnøya. They did not return from one trip to the next to the same foraging area. As the season progressed and the chicks were left alone on the nest, parents increased the duration and maximum range of foraging trips as well as the distance covered. However, they still perform a succession of relatively short foraging trips to the east of the Bjørnøya shelf but they interspersed these short trips with longer foraging trips. One bird returned twice to the same site along the Norwegian coast 570?km from Bjørnøya, the other foraged at 580?km in the mid-Barents Sea. Average flight speed including time spent on the water was 28?km/h and reached 70?km/h during bouts of more than 1?h when the bird was probably continuously in flight.     

Early‐life foraging: Behavioral responses of newly fledged albatrosses to environmental conditions

In order to survive and later recruit into a population, juvenile animals need to acquire resources through the use of innate and/or learnt behaviors in an environment new to them. For far‐ranging marine species, such as the wandering albatross Diomedea exulans, this is particularly challenging as individuals need to be able to rapidly adapt and optimize their movement strategies in response to the highly dynamic and heterogeneous nature of their open‐ocean pelagic habitats. Critical to this is the development and flexibility of dispersal and exploratory behaviors. Here, we examine the movements of eight juvenile wandering albatrosses, tracked using GPS/Argos satellite transmitters for eight months following fledging, and compare these to the trajectories of 17 adults to assess differences and similarities in behavioral strategies through time. Behavioral clustering algorithms (Expectation Maximization binary Clustering) were combined with multinomial regression analyses to investigate changes in behavioral mode probabilities over time, and how these may be influenced by variations in day duration and in biophysical oceanographic conditions. We found that juveniles appeared to quickly acquire the same large‐scale behavioral strategies as those employed by adults, although generally more time was spent resting at night. Moreover, individuals were able to detect and exploit specific oceanographic features in a manner similar to that observed in adults. Together, the results of this study suggest that while shortly after fledging juvenile wandering albatrosses are able to employ similar foraging strategies to those observed in adults, additional skills need to be acquired during the immature period before the efficiency of these behaviors matches that of adults.     

The effects of loggers on the foraging effort and chick-rearing ability of parent little auks

We studied the effects of loggers attached to chick-rearing little auks (Alle alle) on their daily time budget (proportion of time spent in the colony and at sea), foraging activity (duration and proportion of long and short foraging flights), chick provisioning rate and their growth and development on Spitsbergen. We found that experimental parent birds performed shorter but more frequent long foraging flights and reduced the frequency of short foraging flights. They spent more time at the colony and reduced chick provisioning rate compared to control birds. Nestlings reared by experimental parents weighed significantly less at their middle, peak and fledging age and departed colony later than chicks of control parents. Little auks depend on energy-rich copepods associated with cold Arctic waters and are expected to face the climate-induced worsening of the foraging conditions, which may have negative impact on their time/energy budget and survival. The study may help to determine the level of extra effort little auks need to invest to breed successfully.     

Prolonged Parental Feeding in Tool‐Using New Caledonian Crows

According to life‐history theory, the duration of extended parental feeding is determined by the costs and benefits of maximising reproductive success. Therefore, the length of regular parental provisioning should be correlated with the time required for juveniles to acquire the skills that they need to be independent. The relatively few cases of extremely prolonged parental feeding in both land and sea birds appear to be consistent with this prediction because they are associated with learning‐intensive foraging techniques. New Caledonian crows have the most intricate tool manufacture techniques amongst non‐human animals and juveniles take over 1 yr to reach adult‐like proficiency in their tool skills. We investigated the prediction that this species also should have prolonged parental provisioning. We found that these crows have one of the longest known periods of regular extended parental provisioning in birds. Some parents regularly fed juveniles for up to 10 mo post‐fledging. Humans also stand out amongst primates because of their learning‐intensive foraging strategies and an extended period of juvenile dependence. The independently evolved association between a relatively high level of technological skill in foraging and prolonged juvenile provisioning in both humans and New Caledonian crows raises the possibility that these two characteristics might be causally related.     

Wild geese do not increase flight behaviour prior to migration

Hypertrophy of the flight muscles is regularly observed in birds prior to long-distance migrations. We tested the hypothesis that a large migratory bird would increase flight behaviour prior to migration, in order to cause hypertrophy of the flight muscles, and upregulate key components of the aerobic metabolic pathways. Implantable data loggers were used to record year-round heart rate in six wild barnacle geese (Branta leucopsis), and the amount of time spent in flight each day was identified. Time in flight per day did not significantly increase prior to either the spring or the autumn migration, both between time periods prior to migration (5, 10 and 15 days), or when compared with a control period of low activity during winter. The lack of significant increase in flight prior to migration suggests that approximately 22 min per day is sufficient to maintain the flight muscles in condition for prolonged long-distance flight. This apparent lack of a requirement for increased flight activity prior to migration may be attributable to pre-migratory mass gains in the geese increasing workload during short flights, potentially prompting hypertrophy of the flight muscles.     

Gliding flight in the American Kestrel (Falco sparverius): An electromyographic study

Electromyographic (EMG) activity was studied in American Kestrels (Falco sparverius) gliding in a windtunnel tilted to 8 degrees below the horizontal. Muscle activity was observed in Mm. biceps brachii, triceps humeralis, supracoracoideus, and pectoralis, and was absent in M. deltoideus major and M. thoracobrachialis (region of M. pectoralis). These active muscles are believed to function in holding the wing protracted and extended during gliding flight. Quantification of the EMG signals showed a lower level of activity during gliding than during flapping flight, supporting the idea that gliding is a metabolically less expensive form of locomotion than flapping flight. Comparison with the pectoralis musculature of specialized gliding and soaring birds suggests that the deep layer of the pectoralis is indeed used during gliding flight and that the slow tonic fibers found in soaring birds such as vultures represents a specialization for endurant gliding. It is hypothesized that these slow fibers should be present in the wing muscles that these birds use for wing protraction and extension, in addition to the deep layer of the pectoralis. © 1993 Wiley-Liss, Inc.     

Effects of water discharge on fledging time,growth, and survival of piping plovers on the Missouri River

River flow management and modification is a global issue, and its effects on river-dependent organisms are pervasive. Flow modification can directly affect avian species through mortality or habitat loss, but less is known about indirect and sublethal effects of flow modification on reproductive output in these species. Young birds are more vulnerable to predation between hatching and fledging than after flight is achieved, but tradeoffs must be made to balance growth and survival. Predation pressure appears to be a significant factor affecting the time to fledging in altricial birds, but less is known about this threat for precocial birds. Birds reaching fledging earlier should have greater rates of survival to migration because their predator escape repertoire includes flight at an earlier age. We evaluated the effect of varying outflows from the Gavins Point Dam on the growth, age at fledging, and survival of piping plover (Charadrius melodus) chicks on the Missouri River (2006–2009). The study was characterized by 2 relatively high flow years (2006 and 2009) and 2 relatively low flow years (2007 and 2008). We used success rate in recapturing chicks in capture–mark–recapture models as an index for fledging. We attempted to recapture all chicks (n = 1,099) by hand every 3–4 days throughout the season to acquire morphological measurements. Models indicated that as flows from the dam increased, age at fledging increased. We also found that increasing flows were associated with decreasing daily survival rates (β_flow = −2.401, 95% CI: −4.351 to −0.452). Flow was also negatively related to chick mass gain, but we found less evidence for an effect on wing-chord length. Increased flows covered wet-substrate foraging habitat, and likely affected plover reproductive output directly through chick survival and indirectly through decreased growth and increased fledging times. © The Wildlife Society, 2013     

Post-fledging brood and care division in the roseate tern (Sterna dougallii)

Extended post-fledging parental care is an important aspect of parental care in birds, although little studied due to logistic difficulties. Commonly, the brood is split physically (brood division) and/or preferential care is given to a subset of the brood by one parent or the other (care division). Among gulls and tern (Laridae), males and females generally share parental activities during the pre-fledging period, but the allocation of parental care after fledging is little documented. This study examined the behaviour of male and female roseate terns (Sterna dougallii) during the late chick-rearing and early post-fledging periods, and in particular the amount of feeds and the time spent in attendance given to individual chicks/fledglings. Pre-fledging parental care was biparental in all cases. Post-fledging parental care was dependent on the number of fledglings in the brood. Males and females continued biparental care in clutches with one surviving fledgling, while in two-fledgling clutches, males fed the A-fledgling while females fed the B-fledgling. Overall, there was no difference in attendance, only in feeds. This division of care may be influenced by the male only being certain of the paternity of the A-chick but not by chick sex.     

Is mass loss in Brünnich's guillemots Uria lomvia an adaptation for improved flight performance or improved dive performance?

Breeding Brünnich's guillemots Uria lomvia show stepwise mass loss at the time of hatch. This mass loss has usually been explained as an adaptation to reduce the cost of flight during the chick‐rearing period because flight time increases during that period. It is possible, however, that mass loss also increases dive performance during the chick‐rearing period because time spent diving also increases during that period. Reduced mass could reduce basal metabolic rate or costs associated with buoyancy and therefore increase aerobic dive limit. To examine the role of mass loss in dive behavior, we attached time‐depth‐temperature recorders for 24–48 h to chick‐rearing and incubating Brünnich's guillemots at Coats Island, Nunavut (2005: n=45, 2006: n=40), and recorded body mass before and after each deployment. There was no relationship between mass and dive duration during either incubation or chick‐rearing. Seventeen of the birds we sampled during incubation were resampled during chick‐rearing. For this group, dive duration increased with mass loss between incubation and chick‐rearing (r²=0.67–0.75). Mass loss occurred through reductions in metabolically‐active tissues (liver, bladder) and buoyant tissues (lipids) although muscle and gut mass did not change. Despite the large change in lipids, buoyancy only changed by 0.1%, and mass loss therefore did not have much effect on costs associated with buoyancy. Nonetheless, surface pause duration for a given dive depth decreased during chick‐rearing, supporting the idea that reduced mass led to increased aerobic dive limit through reduced metabolic rate and inertial costs; oxygen stores did not increase. We also attached neutrally (n=9) and negatively (n=11) buoyant handicaps to the legs of adults to assess the effect of artificial mass increases on time budgets. Artificially increasing mass decreased total time spent diving but did not change time spent flying. There was no change in shift length between incubation and chick‐rearing, and therefore no support for the idea that mass loss reflected a change in fasting endurance requirements. An energetic model suggested that the observed mass reduction reduced dive costs by 5–8% and flight costs by 3%. We concluded that mass loss may be as important for increasing dive performance as increasing flight performance.     

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 rearing—the most energy-demanding period—and may be related to evolution of RSD.     

Patterns of GPS tracks suggest nocturnal foraging by incubating Peruvian pelicans (Pelecanus thagus)

Most seabirds are diurnal foragers, but some species may also feed at night. In Peruvian pelicans (Pelecanus thagus), the evidence for nocturnal foraging is sparse and anecdotal. We used GPS-dataloggers on five incubating Peruvian pelicans from Isla Lobos de Tierra, Perú, to examine their nocturnality, foraging movements and activities patterns at sea. All instrumented pelicans undertook nocturnal trips during a 5–7 day tracking period. Eighty-seven percent of these trips (n = 13) were strictly nocturnal, whereas the remaining occurred during the day and night. Most birds departed from the island after sunset and returned a few hours after sunrise. Birds traveled south of the island for single-day trips at a maximum range of 82.8 km. Overall, 22% of the tracking period was spent at sea, whereas the remaining time was spent on the island. In the intermediate section of the trip (between inbound and outbound commutes), birds spent 77% of the trip time in floating bouts interspersed by short flying bouts, the former being on average three times longer than the latter. Taken together, the high sinuosity of the bird''s tracks during floating bouts, the exclusively nocturnal trips of most individuals, and the fact that all birds returned to the island within a few hours after sunrise suggest that pelicans were actively feeding at night. The nocturnal foraging strategy of Peruvian pelicans may reduce food competition with the sympatric and strictly diurnal Guanay cormorants (Phalacrocorax bougainvillii), Peruvian boobies (Sula variegata) and Blue-footed boobies (S. nebouxii), which were present on the island in large numbers. Likewise, plankton bioluminescence might be used by pelicans as indirect cues to locate anchovies during their upward migration at night. The foraging success of pelicans at night may be enhanced by seizing prey close to the sea surface using a sit-and-wait strategy.     

Habitat use and movement patterns by dependent and independent juvenile Grasshopper Sparrows during the post‐fledging period

The post‐fledging period is a critical life stage for young grassland birds. Habitat selection by recently fledged birds may differ from that of adults and may change as juveniles transition from the care and protection of parents to independence. To describe patterns of habitat selection during these important life stages, we studied habitat use by juvenile Grasshopper Sparrows (Ammodramus savannarum) in a Conservation Reserve Program grassland in Maryland. We used radio‐telemetry to track daily movement patterns of two age classes of Grasshopper Sparrows during the post‐fledging period. Sparrows were classified as either dependent (<32‐d‐old) or independent (≥32‐d‐old). We characterized the vegetation at 780 vegetation plots (390 plots where birds were located and 390 paired random plots). Microhabitats where dependent birds were found had significantly more bare ground, litter, and plant species richness than paired random plots. In addition, dependent birds were found in plots with less bare ground, more warm‐season grass cover, more total vegetation cover, and more forb cover than plots used by independent birds. Plots where independent birds were located also had significantly more bare ground than random plots. Dependent birds are less able to escape from predators because their flight feathers are not fully grown so they may benefit from remaining in areas of greater vegetation cover. However, juveniles transitioning from dependence to independence must forage on their own, possibly explaining their increased use of more open areas where foraging may be easier. To properly manage habitat for grassland birds, management strategies must consider the changing needs of birds during different stages of development. Our results highlight the importance of diverse grassland ecosystems for juvenile grassland birds during the transition to independence.     

Soaring across continents: decision‐making of a soaring migrant under changing atmospheric conditions along an entire flyway

Thermal soaring birds reduce flight‐energy costs by alternatingly gaining altitude in thermals and gliding across the earth's surface. To find out how soaring migrants adjust their flight behaviour to dynamic atmospheric conditions across entire migration routes, we combined optimal soaring migration theory with high‐resolution GPS tracking data of migrating honey buzzards Pernis apivorus and wind data from a global numerical atmospheric model. We compared measurements of gliding air speeds to predictions based on two distinct behavioural benchmarks for thermal soaring flight. The first being a time‐optimal strategy whereby birds alter their gliding air speeds as a function of climb rates to maximize cross‐country air speed over a full climb– glide cycle (V_opt). The second a risk‐averse energy‐efficient strategy at which birds alter their gliding air speed in response to tailwinds/headwinds to maximize the distance travelled in the intended direction during each glide phase (V_bgw). Honey buzzards were gliding on average 2.05 ms^{– 1} slower than V_opt and 3.42 ms^{– 1} faster than V_bgw while they increased air speeds with climb rates and reduced air speeds in tailwinds. They adopted flexible flight strategies gliding mostly near V_bgw under poor soaring conditions and closer to V_opt in good soaring conditions. Honey buzzards most adopted a time‐optimal strategy when crossing the Sahara, and at the onset of spring migration, where and when they met with the best soaring conditions. The buzzards nevertheless glided slower than V_opt during most of their journeys, probably taking time to navigate, orientate and locate suitable thermals, especially in areas with poor thermal convection. Linking novel tracking techniques with optimal migration models clarifies the way birds balance different tradeoffs during migration.     

Anatomy and histochemistry of spread-wing posture in birds. 3. Immunohistochemistry of flight muscles and the "shoulder lock" in albatrosses

As a postural behavior, gliding and soaring flight in birds requires less energy than flapping flight. Slow tonic and slow twitch muscle fibers are specialized for sustained contraction with high fatigue resistance and are typically found in muscles associated with posture. Albatrosses are the elite of avian gliders; as such, we wanted to learn how their musculoskeletal system enables them to maintain spread-wing posture for prolonged gliding bouts. We used dissection and immunohistochemistry to evaluate muscle function for gliding flight in Laysan and Black-footed albatrosses. Albatrosses possess a locking mechanism at the shoulder composed of a tendinous sheet that extends from origin to insertion throughout the length of the deep layer of the pectoralis muscle. This fascial "strut" passively maintains horizontal wing orientation during gliding and soaring flight. A number of muscles, which likely facilitate gliding posture, are composed exclusively of slow fibers. These include Mm. coracobrachialis cranialis, extensor metacarpi radialis dorsalis, and deep pectoralis. In addition, a number of other muscles, including triceps scapularis, triceps humeralis, supracoracoideus, and extensor metacarpi radialis ventralis, were found to have populations of slow fibers. We believe that this extensive suite of uniformly slow muscles is associated with sustained gliding and is unique to birds that glide and soar for extended periods. These findings suggest that albatrosses utilize a combination of slow muscle fibers and a rigid limiting tendon for maintaining a prolonged, gliding posture.