Vsevolod Afanasyev

Revisiting Levy flight search patterns of wandering albatrosses, bumblebees and deer

The study of animal foraging behaviour is of practical ecological importance, and exemplifies the wider scientific problem of optimizing search strategies. Lévy flights are random walks, the step lengths of which come from probability distributions with heavy power-law tails, such that clusters of short steps are connected by rare long steps. Lévy flights display fractal properties, have no typical scale, and occur in physical and chemical systems. An attempt to demonstrate their existence in a natural biological system presented evidence that wandering albatrosses perform Lévy flights when searching for prey on the ocean surface. This well known finding was followed by similar inferences about the search strategies of deer and bumblebees. These pioneering studies have triggered much theoretical work in physics (for example, refs 11, 12), as well as empirical ecological analyses regarding reindeer, microzooplankton, grey seals, spider monkeys and fishing boats. Here we analyse a new, high-resolution data set of wandering albatross flights, and find no evidence for Lévy flight behaviour. Instead we find that flight times are gamma distributed, with an exponential decay for the longest flights. We re-analyse the original albatross data using additional information, and conclude that the extremely long flights, essential for demonstrating Lévy flight behaviour, were spurious. Furthermore, we propose a widely applicable method to test for power-law distributions using likelihood and Akaike weights. We apply this to the four original deer and bumblebee data sets, finding that none exhibits evidence of Lévy flights, and that the original graphical approach is insufficient. Such a graphical approach has been adopted to conclude Lévy flight movement for other organisms, and to propose Lévy flight analysis as a potential real-time ecosystem monitoring tool. Our results question the strength of the empirical evidence for biological Lévy flights.

Tracking Long-Distance Songbird Migration by Using Geolocators

We mapped migration routes of migratory songbirds to the Neotropics by using light-level geolocators mounted on breeding purple martins (Progne subis) and wood thrushes (Hylocichla mustelina). Wood thrushes from the same breeding population occupied winter territories within a narrow east-west band in Central America, suggesting high connectivity of breeding and wintering populations. Pace of spring migration was rapid (233 to 577 kilometers/day) except for one individual (159 kilometers/day) who took an overland route instead of crossing the Gulf of Mexico. Identifying songbird wintering areas and migration routes is critical for predicting demographic consequences of habitat loss and climate change in tropical regions.

SUMMER DISTRIBUTION AND MIGRATION OF NONBREEDING ALBATROSSES: INDIVIDUAL CONSISTENCIES AND IMPLICATIONS FOR CONSERVATION

Many birds show a surprising degree of intraspecific variability in migratory tendency and choice of wintering site. In this study, we tracked the seasonal movements of 35 nonbreeding Black-browed Albatrosses Thalassarche melanophrys from South Georgia, including 24 birds followed in two consecutive years. This revealed consistent patterns of status-related, sex-specific, and individual variation in wintering strategies, and provided the first description of the summer distribution of failed/deferring breeders. Individuals exhibited a striking degree of site fidelity, returning to the same region (southwest Africa or Australia) and showing correlated centers of distribution, as well as remarkable consistency in the chronology of their movements, in consecutive years. Nonetheless, a degree of behavioral flexibility remained, and particularly on the return migration, birds moved between, or bypassed, alternative intermediate staging sites depending on local circumstances. Initiation of the outward migration varied according to breeding status, timing of failure, and sex: deferring breeders and those that failed early departed two months before successful birds, and successful females departed 1–2 weeks earlier than males. Sex-related latitudinal variation in distribution was also apparent, with females wintering farther north within the Benguela system. Moreover, the only migrant to Australia was a male, supporting an apparent tendency for male-biased breeding dispersal inferred from genetic analyses. Distribution and timing of movements appeared in general to relate to avoidance of competition from congeners and conspecifics from other populations. From a conservation perspective, the study indicated that, for the declining Black-browed Albatross population at South Georgia, the primary focus should be toward improving the management (especially reducing bycatch levels) of fisheries in the central and eastern South Atlantic.

The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration

The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d–1) than during spring (233 km d–1). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.

Year-Round Tracking of Small Trans-Saharan Migrants Using Light-Level Geolocators

Since 1899 ringing (or banding) remained the most important source of information about migration routes, stopover sites and wintering grounds for birds that are too small to carry satellite-based tracking systems. Despite the large quantity of migrating birds ringed in their breeding areas in Europe, the number of ring recoveries from sub-Saharan Africa is very low and therefore the whereabouts of most small bird species outside the breeding season remain a mystery. With new miniaturized light-level geolocators it is now possible to look beyond the limits of ring recovery data. Here we show for the first time year round tracks of a near passerine trans-Saharan migrant, the European Hoopoe (Upupa epops epops). Three birds wintered in the Sahel zone of Western Africa where they remained stationary for most of the time. One bird chose a south-easterly route following the Italian peninsula. Birds from the same breeding population used different migration routes and wintering sites, suggesting a low level of migratory connectivity between breeding and wintering areas. Our tracking of a near passerine bird, the European Hoopoe, with light-level geolocators opens a new chapter in the research of Palaearctic-African bird migration as this new tool revolutionizes our ability to discover migration routes, stopover sites and wintering grounds of small birds.

Anomalous fluctuations in the dynamics of complex systems: from DNA and physiology to econophysics

We discuss examples of complex systems composed of many interacting subsystems. We focus on those systems displaying nontrivial long-range correlations. These include the one-dimensional sequence of base pairs in DNA, the sequence of flight times of the large seabird Wandering Albatross, and the annual fluctuations in the growth rate of business firms. We review formal analogies in the models that describe the observed long-range correlations, and conclude by discussing the possibility that behavior of large numbers of humans (as measured, e.g., by economic indices) might conform to analogs of the scaling laws that have proved useful in describing systems composed of large numbers of inanimate objects.

Lévy flights in random searches

We review the general search problem of how to find randomly located objects that can only be detected in the limited vicinity of a forager, and discuss its quantitative description using the theory of random walks. We illustrate Levy flight foraging by comparison to Brownian random walks and discuss experimental observations of Levy flights in biological foraging. We review recent findings suggesting that an inverse square probability density distribution P(l)∼l−2 of step lengths l can lead to optimal searches. Finally, we survey the explanations put forth to account for these unexpected findings.

Extrinsic and intrinsic determinants of winter foraging and breeding phenology in a temperate seabird

In temperate regions, winter presents animals with a number of challenges including depressed food abundance, increased daily energy requirements, higher frequency of extreme weather events and shortened day length. Overcoming these constraints is critical for overwintering survival and scheduling of future breeding of long-lived species and is likely to be state dependent, associated with intrinsic abilities such as food acquisition rates. We examined the relationship between environmental and intrinsic factors on overwintering foraging and subsequent breeding phenology of the European shag Phalacrocorax aristotelis, a diurnal marine predator. We tested a range of hypotheses relating to overwintering foraging time and location. We found that individuals greatly increased their foraging time in winter to a peak of more than 90% of available daylight at the winter solstice. The seasonal patterns of foraging time appear to be driven by a combination of light levels and weather conditions and may be linked to the availability of the shags principal prey, the lesser sandeel Ammodytes marinus. There was no evidence that shags dispersed south in winter to increase potential foraging time. Foraging time decreased after the winter solstice and, crucially, was correlated with subsequent breeding phenology, such that individuals that spent less time foraging in February bred earlier. The relationship was much stronger in females than males, in line with their more direct control of timing of breeding. Our results demonstrate that pre-breeding intrinsic foraging ability is critical in determining breeding phenology.

Effects of post-breeding moult and energetic condition on timing of songbird migration into the tropics

Each autumn billions of songbirds migrate between the temperate zone and tropics, but little is known about how events on the breeding grounds affect migration to the tropics. Here, we use light level geolocators to track the autumn migration of wood thrushes Hylocichla mustelina and test for the first time if late moult and poor physiological condition prior to migration delays arrival on the winter territory. Late nesting thrushes postponed feather moult, and birds with less advanced moult in August were significantly farther north on 10 October while en route to the tropics. Individuals in relatively poor energetic condition in August (high β-Hydroxybutyrate, low triglyceride, narrow feather growth bars) passed into the tropics significantly later in October. However, late moult and poor pre-migratory condition did not result in late arrival on the winter territory because stopover duration was highly variable late in migration. Although carry-over effects from the winter territory to spring migration may be strong in migratory songbirds, our study suggests that high reproductive effort late in the season does not impose time constraints that delay winter territory acquisition.

From cradle to early grave: juvenile mortality in European shags Phalacrocorax aristotelis results from inadequate development of foraging proficiency

In most long-lived animal species, juveniles survive less well than adults. A potential mechanism is inferior foraging skills but longitudinal studies that follow the development of juvenile foraging are needed to test this. We used miniaturized activity loggers to record daily foraging times of juvenile and adult European shags Phalacrocorax aristotelis from fledging to the following spring. Juveniles became independent from their parents 40 days post-fledging. They compensated for poor foraging proficiency by foraging for approximately 3 h d−1 longer than adults until constrained by day length in early November. Thereafter, juvenile foraging time tracked shortening day length up to the winter solstice, when foraging time of the two age classes converged and continued to track day length until early February. Few individuals died until midwinter and mortality peaked in January–February, with juvenile mortality (including some of the study birds) five times that of adults. In their last two weeks of life, juveniles showed a marked decline in foraging time consistent with individuals becoming moribund. Our results provide compelling evidence that juveniles compensate for poor foraging proficiency by increasing foraging time, a strategy that is limited by day length resulting in high winter mortality.