Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

Flight distance of urban birds, predation, and selection for urban life

Numerous species have adapted to humans, especially invasive species associated with humans in towns and cities. Short flight distances of populations adapted to urban environments reflect changes in behavior and physiology, reflecting phenotypic plasticity or evolution. Here, I tested the hypothesis that the decrease in flight distance to a potential predator (an approaching human) reflected adaptation to urbanization, using a data set of flight distances of 44 common species of European birds in different stages of adaptation to urban environments. Urban populations had consistently shorter flight distances than rural populations of the same species. Variation in relative flight distance of urban populations was predicted by the number of generations since urbanization, as expected by a gradual process of adaptation. Furthermore, species with relatively large populations in urban environments would be an indication of local adaptation to urban environments. Relative flight distance of urban population was shorter for species with large populations in urban compared to rural habitats. Species that had adapted to urban environments as shown by short flight distances were less susceptible to predation by the European sparrowhawk Accipiter nisus than species with relatively long flight distances in urban populations. These findings provide evidence consistent with the hypothesis that recent changes in the tameness of urban birds, as reflected by their relatively short flight distances, is an adaptation to the novel urban environment. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Timing of arrival from spring migration is associated with flight performance in the migratory barn swallow

Timing of arrival at the breeding grounds by migratory birds affects their mating success and access to superior resources, thus being a major factor associated with fitness. Much empirical work has been devoted to investigate the condition dependence of arrival sequence of migrants and characteristics of individuals that influence arrival time from migration. Surprisingly, there are no studies examining the relationship between flight performance of individual birds and their arrival time. I investigated the relative importance of direct effects of short-term flight performance, age, body condition and the degree of sexual ornamentation (tail length) on timing of spring arrival in the barn swallow (Hirundo rustica), a long-distance trans-equatorial passerine migrant. I evaluated short-term flight performance (a composite variable comprising flight manoeuvrability, velocity and acceleration) in a standardised manner using flight tunnels. Short-term flight performance was a significant and important predictor of spring arrival date. Furthermore, locomotion predicted arrival date of individual birds independently of morphological variables—the degree of sexual ornamentation (the length of the tail) and wing aspect ratio and body condition. I discuss the possible role short-term flight performance may have in determining migratory performance. This is the first time flight performance has been shown to be associated with timing of arrival from migration in a migratory bird.     

Evidence for two genetically and chemically defined host races of Tyria jacobaeae (Arctiidae, Lepidoptera)

Summary. A widely distributed host race of Tyria jacobaeae lives on Senecio jacobaea and related species and accumulates pyrrolizidine alkaloids (“PA race”), another race, which is restricted to the Alps and found on Petasites paradoxus, sequesters sesquiterpenes, such as petasol and isopetasol. Nucleotide sequences of the mitochondrial 16S rDNA gene show 1% sequence divergence, indicating that genetical differences exist between the PA exploiting and the terpene-sequestering host races of T. jacobaeae. This finding suggests that both host races of T. jacobaeae must have been separated for some time already, possibly since the Pleistocene. Received 2 May 2001; accepted 1 June 2001.     

Costs of overlapping reproduction and moult in passerine birds: an experiment with the pied flycatcher

The process of moult in birds requires the investment of substantial amounts of energy and nutrients in feather production and attendant processes. Flight performance may be reduced during the moult of the wing feathers, and moulting birds may suffer from an increased predation risk. These factors may explain why the moult is usually separated in time from other energetically demanding processes, such as reproduction. In this study, we investigated the importance of the temporal separation of moult from breeding activities in the pied flycatcher Ficedula hypoleuca. We induced a moult-breeding overlap by removing the two innermost primaries on both wings, thus imitating the natural loss of these feathers during the initial stage of the moult. The experiment probably did not stimulate feather regrowth, but may have reduced flight performance. Just before fledging of the young, manipulated males and females, as well as the fledglings, had a lower body condition than control birds. The return rate of adult males in the year following the manipulation was significantly lower for males in the manipulated group than for control males. The return rate of females was lower than that of males, but there was no difference between the two female groups. The number of nestlings recruited in the year after the manipulation was significantly lower for the group with an induced moult-breeding overlap than for the control group. The reproductive performance in recruited and returning old females was independent of the manipulation, and in returning adults and recruits the feather quality in terms of number of barbs/mm was independent of the manipulation. These results suggest that a moult-breeding overlap induces fitness costs in terms of reduced recruitment rate and adult survival in a long-distance migrant. Received: 17 September 1997 / Accepted after revision: 23 February 1998     

Transoceanic bird migration: Evidence for detection of wind direction

Summary Further analysis of radar tracks of migrating birds over the Atlantic Ocean (Larkin et al., 1979) indicates that birds sometimes fly at lower air speeds when winds are in the direction of flight than in crosswinds or head winds. The birds are able to estimate wind direction over open ocean by a mechanism which is presently open to speculation.     

Long-range migratory travel of a green turtle tracked by satellite: evidence for navigational ability in the open sea

A female of Chelonia mydas was tracked by satellite in the South China Sea in 1993 from the nesting beach to the resident foraging grounds more than 600 km away. The final leg of the journey, 475 km long, directly pinpointed the goal, with the turtle maintaining a constant speed and direction both night and day. This provides clues about the navigational mechanism used.     

Obligatory barrier crossing and adaptive fuel management in migratory birds: the case of the Atlantic crossing in Northern Wheatears (<Emphasis Type="Italic">Oenanthe oenanthe</Emphasis>)

Behaviour on migration was often suggested to be selected for time-minimising strategies. Current optimality models predict that optimal fuel loads at departure from stopover sites should increase with increasing fuel deposition rates. We modified such models for the special case of the east Atlantic crossing of the Northern Wheatear (Oenanthe oenanthe). From optimality theory, we predict that optimal time-minimising behaviour in front of such a barrier should result in a positive correlation between fuel deposition rates and departure fuel loads only above a certain threshold, which is the minimum fuel load (f _min) required for the barrier crossing. Using a robust range equation, we calculated the minimum fuel loads for different barrier crossings and predict that time-minimising wheatears should deposit a minimum of 24% fuel in relation to lean body mass (m ₀ ) for the sea crossing between Iceland and Scotland. Fuel loads of departing birds in autumn in Iceland reached this value only marginally but showed positive correlation between fuel deposition rate (FDR) and departure fuel load (DFL). Birds at Fair Isle (Scotland) in spring, which were heading towards Iceland or Greenland, were significantly heavier and even showed signs of overloading with fuel loads up to 50% of lean body mass. Departure decisions of Icelandic birds correlated significantly with favourable wind situations when assuming a migration direction towards Spain; however, the low departure fuel loads contradict a direct non-stop flight.     

A case for quantile regression in behavioral ecology: getting more out of flight initiation distance data

A central goal of behavioral ecology is to quantify and explain variation in behavior. While much previous work has focused on the differences in mean behavior across groups or treatments, we present a complementary approach studying changes in the distribution of the response variable. This is important because changes in the edges of a distribution may be more informative than changes in the mean if behavior at the edges of a distribution better reflects behavioral constraints. Quantile regression estimates the rate of change of conditional quantiles of a response variable and thus allows the study of changes in any part of its distribution. Although quantile regression is gaining popularity in the ecological literature, it is strikingly unused in behavioral ecology. Here, we demonstrate the usefulness of this method by analyzing the relationship between the starting distance (SD) at which an observer approach a focal animal and its flight initiation distance (FID, the distance between the observer and the animal when it decides to flee). In particular, we used a simple model of flight initiation distance to show that in most situations ordinary least-square regression cannot be used to analyse the SD–FID relationship. Quantile regression conducted on the lowest quantiles appears more robust and we applied this approach to data from four bird species. Overall, changes in the lowest FID values appeared to be the most informative to determine if a species displays a “flush early” strategy, a strategy which has been hypothesized to be a general rule. We hope this example will bring quantile regression to the attention of behavioral ecologists as a valuable tool to add to their statistical toolbox.     

Inheritance of migratory direction in a bird species: a cross-breeding experiment with SE- and SW-migrating blackcaps (Sylvia atricapilla)

Summary Young avian migrants of many species are able to find their species- or population-specific wintering area without the help of conspecifics. In orientation tests hand-raised birds have been demonstrated to choose appropriate population-specific migratory directions, suggesting a genetic basis to this behaviour. I here report results of a cross-breeding experiment between individuals of two blackcap (Sylvia atricapilla) populations with widely different migratory directions. The orientation of the F1 offspring was intermediate between and significantly different from that of both parental populations (Fig. 2). The variance of individual mean directions in the F1 generation did not increase compared with the parental groups, and the inheritance of migratory directions was not sex-linked. The data provide direct evidence for a genetic basis of migratory directions in birds and demonstrate a phenotypically intermediate mode of inheritance.     

Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection

We studied the echolocation and hunting behavior of three aerial insectivorous species of bats (Vespertilionidae: Pipistrellus) in the field in order to characterize the signals used by the bats and to determine how call structure varies in relation to habitat structure (uncluttered versus cluttered space). We documented free-flying, naturally foraging wild pipistrelles in various habitats using multiflash stereophotography combined with simultaneous sound recordings. Then we reconstructed the bat's flight position in three-dimensional space and correlated it with the corresponding echolocation sequences. In all three species of pipistrelles, signal structure varied substantially. In echolocation sequences of the search phase we found a consistent association of signal types with habitat types. In uncluttered habitats (obstacles more than 5 m from the bat) pipistrelles emitted almost exclusively narrowband signals with bandwidths less than 15 kHz. In cluttered habitats (obstacles less than 5 m from the bat) they switched to signals with bandwidths of more than 15 kHz. Wideband signals were also used when the bats were turning in cluttered and uncluttered spaces and for an instant after turning away from obstacles. Prey detection occured only when the outgoing signal did not overlap with the returning echo from potential prey. The bats also avoided overlap of echoes from potential prey and obstacles. Based on the results of this study, we propose an overlap-free window within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects. Correspondence to: E.K.V. Kalko     

Mechanisms of crustacean egg hatching: Evidence for enzyme release by crab embryos

During the summer of 1989, we examined mechanisms of egg hatching in three species of brachyurans that occupy different habitats as adults near Beaufort, North Carolina, USA:Neopanope sayi (Smith) (subtidal),Uca pugilator (Bosc) (intertidal) andSesarma cinereum (Bosc) (supratidal). Results of casein assays indicated that embryos of all species release proteolytic enzymes near the time of egg hatching. Species differences in specific enzyme activity were suggested, with increased activity in the more terrestrial crabs. Embryos ofN. sayi released enzymes several hours before larval release by the female, whileU. pugilator andS. cinereum released enzymes closer to the time of larval release; enzyme release coincided roughly with time of egg-membrane breakage in all species. Direct observations of hatching showed a sequence of outer-membrane breakage apparently followed by inner-membrane breakage and emergence of the larva. Egg volumes increased most during early and/or mid-stages of development, with a marked slowing of the increase during the several days before hatching. Thus, a gradual osmotically-driven increase in water content may also be involved in egg hatching.     

Optimal Institutional Arrangements for Transboundary Pollutants in a Second-Best World: Evidence from a Differential Game with Asymmetric Players

This paper uses a dynamic model with asymmetric players to explore the question: In a second-best world, should environmental regulations for transboundary pollutants be carried out locally or centrally? We find that combined payoffs are larger with decentralized control if payoffs are sufficiently heterogeneous and initial pollution stocks are sufficiently small. This result is obtained because the central authority applies one shadow price to pollution (i.e., it uses uniform standards), whereas local authorities use different shadow prices, and therefore different standards.     

Maternal energy expenditure does not change with flight costs or food availability in the pied flycatcher (Ficedula hypoleuca): costs and benefits for nestlings

We manipulated parental work load without changing brood size in a population of pied flycatchers Ficedula hypoleuca by removing two primaries (7 and 9) from each wing of females, thus reducing wing area and increasing flight costs. At other nests, we offered supplementary food in the form of live mealworms (10–20 g daily from hatching) to reduce brood demand and thus parental foraging costs. Other nests were left as controls. The daily energy expenditure of females feeding 12-day-old nestlings was measured with doubly labelled water D₂ ¹⁸O. Females in both treatments expended the same amount of energy, fed at the same rate and had similar body masses to birds in the control group. No effect of treatment on male mass and feeding effort was detected. More nestlings, however, died in nests of handicapped females. Nestlings of handicapped females had significantly lower body mass and haematocrit values than nestlings in food-supplemented nests, with nestlings in control nests occupying an intermediate position. The effects of both treatments on nestling mass, haematocrit values and mortality rates were only noticeable in nests infested with mites. Maternal energy expenditure is apparently constrained and offspring pay the costs imposed by reduced provisioning rate or increased demand caused by ectoparasites, while receiving benefits when food supply improves. The presumption that avian reproductive costs derive from changes in a flexible energy output may not be met in many cases. Received: 24 October 1998 / Received in revised form: 15 March 1999 / Accepted: 26 April 1999     

Effects of experimental protocol on global vegetation model accuracy: A comparison of simulated and observed vegetation patterns for Asia

Prognostic vegetation models have been widely used to study the interactions between environmental change and biological systems. This study examines the sensitivity of vegetation model simulations to: (i) the selection of input climatologies representing different time periods and their associated atmospheric CO₂ concentrations, (ii) the choice of observed vegetation data for evaluating the model results, and (iii) the methods used to compare simulated and observed vegetation. We use vegetation simulated for Asia by the equilibrium vegetation model BIOME4 as a typical example of vegetation model output. BIOME4 was run using 19 different climatologies and their associated atmospheric CO₂ concentrations. The Kappa statistic, Fuzzy Kappa statistic and a newly developed map-comparison method, the Nomad index, were used to quantify the agreement between the biomes simulated under each scenario and the observed vegetation from three different global land- and tree-cover data sets: the global Potential Natural Vegetation data set (PNV), the Global Land Cover Characteristics data set (GLCC), and the Global Land Cover Facility data set (GLCF). The results indicate that the 30-year mean climatology (and its associated atmospheric CO₂ concentration) for the time period immediately preceding the collection date of the observed vegetation data produce the most accurate vegetation simulations when compared with all three observed vegetation data sets. The study also indicates that the BIOME4-simulated vegetation for Asia more closely matches the PNV data than the other two observed vegetation data sets. Given the same observed data, the accuracy assessments of the BIOME4 simulations made using the Kappa, Fuzzy Kappa and Nomad index map-comparison methods agree well when the compared vegetation types consist of a large number of spatially continuous grid cells. The results of this analysis can assist model users in designing experimental protocols for simulating vegetation.     

Waterfowls habitat modeling: Simulation of nest site selection for the migratory Little Tern (Sterna albifrons) in the Nakdong estuary

This paper aims to find patterns in nest site selection by Little Terns Sterna albifrons, in the Nakdong estuary in South Korea. This estuary is important waterfowl stopover and breeding habitat, located in the middle of the East Asia-Australasian Flyway. The Little Tern is a common species easily observed near the seashore but their number is gradually declining around the world. We investigated their nests and eggs on a barrier islet in the Nakdong estuary during the breeding season (May to June, 2007), and a pattern for the nest site selection was identified using genetic programming (GP). The GP generated a predictive rule-set model for the number of Little Tern nests (training: R² = 0.48 and test: 0.46). The physical features of average elevation, variation of elevation, plant coverage, and average plant height were estimated to determine the influence on nest numbers for Little Tern. A series of sensitivity analyses stressed that mean elevation and vegetation played an important role in nest distribution for Little Tern. The influence of these two variables could be maximized when elevation changed moderately within the sampled quadrats. The study results are regarded as a good example of applying GP to vertebrate distribution patterning and prediction with several important advantages compared to conventional modeling techniques, and can help establish a management or restoration strategy for the species.     

Innate movement rules in foraging bees: flight distances are affected by recent rewards and are correlated with choice of flower type

The non-random movement patterns of foraging bees are believed to increase their search efficiency. These patterns may be innate, or they may be learned through the bees’ early foraging experience. To identify the innate components of foraging rules, we characterized the flight of naive bumblebees, foraging on a non-patchy “field” of randomly scattered artificial flowers with three color displays. The flowers were randomly mixed and all three flower types offered equal nectar volumes. Visited flowers were refilled with probability 0.5. Flight distances, flight durations and nectar probing durations were determined and related to the bees’ recent experiences. The naive bees exhibited area-restricted search behavior, i.e., flew shorter distances following visits to rewarding flowers than after visits to empty flowers. Additionally, flight distances during flower-type transitions were longer than flight distances between flowers of the same type. The two movement rules operated together: flight distances were longest for flights between flower types following non-rewarding visits, shortest for within-type flights following rewarding visits. An increase in flight displacement during flower-type shifts was also observed in a second experiment, in which all three types were always rewarding. In this experiment, flower-type shifts were also accompanied by an increase in flight duration. Possible relationships between flight distances, flight durations and flower-type choice are discussed. Received: 20 November 1995/Accepted after revision: 10 May 1996