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Terminal attack trajectories of peregrine falcons are described by the proportional navigation guidance law of missiles.

ABSTRACT: The ability to intercept uncooperative targets is key to many diverse flight behaviors, from courtship to predation. Previous research has looked for simple geometric rules describing the attack trajectories of animals, but the underlying feedback laws have remained obscure. Here, we use GPS loggers and onboard video cameras to study peregrine falcons, Falco peregrinus, attacking stationary targets, maneuvering targets, and live prey. We show that the terminal attack trajectories of peregrines are not described by any simple geometric rule as previously claimed, and instead use system identification techniques to fit a phenomenological model of the dynamical system generating the observed trajectories. We find that these trajectories are best-and exceedingly well-modeled by the proportional navigation (PN) guidance law used by most guided missiles. Under this guidance law, turning is commanded at a rate proportional to the angular rate of the line-of-sight between the attacker and its target, with a constant of proportionality (i.e., feedback gain) called the navigation constant (N). Whereas most guided missiles use navigation constants falling on the interval 3 ? N ? 5, peregrine attack trajectories are best fitted by lower navigation constants (median N < 3). This lower feedback gain is appropriate at the lower flight speed of a biological system, given its presumably higher error and longer delay. This same guidance law could find use in small visually guided drones designed to remove other drones from protected airspace.

SUBMITTER: Brighton CH

PROVIDER: S-EPMC5754800 | biostudies-literature | 2017 Dec

REPOSITORIES: biostudies-literature

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Terminal attack trajectories of peregrine falcons are described by the proportional navigation guidance law of missiles.

Brighton Caroline H CH Thomas Adrian L R ALR Taylor Graham K GK

Proceedings of the National Academy of Sciences of the United States of America 20171204 51

The ability to intercept uncooperative targets is key to many diverse flight behaviors, from courtship to predation. Previous research has looked for simple geometric rules describing the attack trajectories of animals, but the underlying feedback laws have remained obscure. Here, we use GPS loggers and onboard video cameras to study peregrine falcons, <i>Falco peregrinus</i>, attacking stationary targets, maneuvering targets, and live prey. We show that the terminal attack trajectories of pereg ...[more]

PMID: 29203660

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Project description:Subspecies relationships within the peregrine falcon (Falco peregrinus) have been long debated because of the polytypic nature of melanin-based plumage characteristics used in subspecies designations and potential differentiation of local subpopulations due to philopatry. In North America, understanding the evolutionary relationships among subspecies may have been further complicated by the introduction of captive bred peregrines originating from non-native stock, as part of recovery efforts associated with mid 20th century population declines resulting from organochloride pollution. Alaska hosts all three nominal subspecies of North American peregrine falcons-F. p. tundrius, anatum, and pealei-for which distributions in Alaska are broadly associated with nesting locales within Arctic, boreal, and south coastal maritime habitats, respectively. Unlike elsewhere, populations of peregrine falcon in Alaska were not augmented by captive-bred birds during the late 20th century recovery efforts. Population genetic differentiation analyses of peregrine populations in Alaska, based on sequence data from the mitochondrial DNA control region and fragment data from microsatellite loci, failed to uncover genetic distinction between populations of peregrines occupying Arctic and boreal Alaskan locales. However, the maritime subspecies, pealei, was genetically differentiated from Arctic and boreal populations, and substructured into eastern and western populations. Levels of interpopulational gene flow between anatum and tundrius were generally higher than between pealei and either anatum or tundrius. Estimates based on both marker types revealed gene flow between augmented Canadian populations and unaugmented Alaskan populations. While we make no attempt at formal taxonomic revision, our data suggest that peregrine falcons occupying habitats in Alaska and the North Pacific coast of North America belong to two distinct regional groupings-a coastal grouping (pealei) and a boreal/Arctic grouping (currently anatum and tundrius)-each comprised of discrete populations that are variously intra-regionally connected.

Project description:High levels (microg/g lw) of polybrominated diphenyl ethers (PBDEs) and polychlorinated biphenyls (PCBs) were measured in peregrine falcon eggs from California (n = 90 eggs from 52 birds, 38 nest sites, collected 1986-2007, SigmaPBDEs median = 4.53, range = 0.08-53.1). Over the past 22 years, PBDE levels more than tripled each decade in the eggs, whereas PCB levels had no significant changes. PBDE levels were highest in eggs from major California cities ("Big Cities"), whereas PCBs showed no difference across the regions. For PBDEs, Big City eggs had markedly different patterns from Coastal eggs: BDE-209 and the higher brominated PBDEs (hexa-nona) were dominant congeners in Big City eggs, while BDE-47 and -99 were dominant in Coastal eggs. In many of the birds that gave multiple eggs over time ("time series"), PBDE patterns changed over time: the high proportions of BDE-209 and higher brominated PBDEs (short half-lives) in young birds contrasted with increasingly higher proportions of BDE-153 (long half-life) and other lower brominated PBDEs as the birds aged. These data are consistent with metabolic debromination of BDE-209 (t(1/2) = 1-2 weeks) to the lower brominated PBDEs, with accumulation over time of BDE-153 (t(1/2) = 3-4 years). In contrast, PCB patterns showed no differences by locations, and did not change over time. Diet (prey birds) may explain the urban PBDE pattern, as the patterns in urban pigeons and peregrines were similar, with high proportions of BDE-209 and the higher-brominated PBDEs. Also, our prey data (feathers from peregrine nests) showed urban peregrines having a higher proportion (>2 fold) of granivorous/opportunistic birds (e.g., "introduced feral" pigeons, mourning doves, starlings) in their diet than coastal peregrines. In summary, these data indicate that BDE-209 exits consumer products as an environmental contaminant to be taken up by wildlife (particularly in urban locations), and undergoes metabolic debromination to the banned lower-brominated PBDEs. High levels of the higher-brominated PBDE congeners, especially in urban locations, permitted accurate measures of relative proportions of homologues in each of the hexa-nona congener classes. Using the major hexa-nona homologues in each of these classes, we propose a pathway for the stepwise, metabolic debromination of BDE-209.

Project description:BackgroundOur ability to monitor populations or species that were once threatened or endangered and in the process of recovery is enhanced by using genetic methods to assess overall population stability and size over time. This can be accomplished most directly by obtaining genetic measures from temporally-spaced samples that reflect the overall stability of the population as given by changes in genetic diversity levels (allelic richness and heterozygosity), degree of population differentiation (F(ST) and D(EST)), and effective population size (N(e)). The primary goal of any recovery effort is to produce a long-term self-sustaining population, and these genetic measures provide a metric by which we can gauge our progress and help make important management decisions.Methodology/principal findingsThe peregrine falcon in North America (Falco peregrinus tundrius and anatum) was delisted in 1994 and 1999, respectively, and its abundance will be monitored by the species Recovery Team every three years until 2015. Although the United States Fish and Wildlife Service makes a distinction between tundrius and anatum subspecies, our genetic results based on eleven microsatellite loci suggest limited differentiation that can be attributed to an isolation by distance relationship and warrant no delineation of these two subspecies in its northern latitudinal distribution from Alaska through Canada into Greenland. Using temporal samples collected at Padre Island, Texas during migration (seven temporal time periods between 1985-2007), no significant differences in genetic diversity or significant population differentiation in allele frequencies between time periods were observed and were indistinguishable from those obtained from tundrius/anatum breeding locations throughout their northern distribution. Estimates of harmonic mean N(e) were variable and imprecise, but always greater than 500 when employing multiple temporal genetic methods.Conclusions/significanceThese results, including those from simulations to assess the power of each method to estimate N(e), suggest a stable or growing population, which is consistent with ongoing field-based monitoring surveys. Therefore, historic and continuing efforts to prevent the extinction of the peregrine falcon in North America appear successful with no indication of recent decline, at least from the northern latitude range-wide perspective. The results also further highlight the importance of archiving samples and their use for continual assessment of population recovery and long-term viability.

Project description:When aiming to capture a fast-moving target, animals can follow it until they catch up, or try to intercept it. In principle, interception is the more complicated strategy, but also more energy efficient. To study whether simple feedback controllers can explain interception behaviours by animals with miniature brains, we have reconstructed and studied the predatory flights of the robber fly Holcocephala fusca and killer fly Coenosia attenuata Although both species catch other aerial arthropods out of the air, Holcocephala contrasts prey against the open sky, while Coenosia hunts against clutter and at much closer range. Thus, their solutions to this target catching task may differ significantly. We reconstructed in three dimensions the flight trajectories of these two species and those of the presented targets they were attempting to intercept. We then tested their recorded performances against simulations. We found that both species intercept targets on near time-optimal courses. To investigate the guidance laws that could underlie this behaviour, we tested three alternative control systems (pure pursuit, deviated pursuit and proportional navigation). Only proportional navigation explains the timing and magnitude of fly steering responses, but with differing gain constants and delays for each fly species. Holcocephala uses a dimensionless navigational constant of N ≈ 3 with a time delay of ≈28 ms to intercept targets over a comparatively long range. This constant is optimal, as it minimizes the control effort required to hit the target. In contrast, Coenosia uses a constant of N ≈ 1.5 with a time delay of ≈18 ms, this setting may allow Coenosia to cope with the extremely high line-of-sight rotation rates, which are due to close target proximity, and thus prevent overcompensation of steering. This is the first clear evidence of interception supported by proportional navigation in insects. This work also demonstrates how by setting different gains and delays, the same simple feedback controller can yield the necessary performance in two different environments.

Dataset Information

Terminal attack trajectories of peregrine falcons are described by the proportional navigation guidance law of missiles.

Publications

Terminal attack trajectories of peregrine falcons are described by the proportional navigation guidance law of missiles.

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