Effects of boundary layer conductance on substomatal pressures of carbon dioxide

Abstract. Gas exchange measurements were made on single leaves of three C₃ and one C₄ species at air speeds of 0.4 and 4.0 m s⁻¹ to determine if boundary layer conductance substantially affected the substomatal pressure of carbon dioxide. Boundary layer conductances to water vapour were 0.4 to 0.5 mol m⁻² s⁻¹ at the lower air speed, and 1.2 to 1.5 mol m⁻² s⁻¹ at the higher air speed. Substomatal carbon dioxide pressures were about 5 Pa lower at low boundary layer conductance in the C₃ species, and about 3 Pa lower in the C₄ species when measurements were made at high and moderate photosynthetic photon flux densities. No evidence of stomatal adjustment to altered boundary layer conductance was found. Photosynthetic rates at high photon flux densities were reduced by about 20% at the low air speed in the C₃ species. The commonly reported values of substomatal carbon dioxide pressure for C₃ and C₄ species were found to occur only when measurements were made at the higher air speed.     

How swift are swifts Apus apus?

Swifts Apus apus are renowned for their fast flight manner which has fascinated people in all times. However, previous studies of swifts in flight during migration and roosting flights have shown that the birds operate over a narrow range of flight speeds compared with most other birds studied. In this study we have focused on the special flight behavior often called 'screaming parties'. During these flights the birds appear to reach very high speeds and therefore we used a stereo high speed camera setup to measure the flight speeds of the birds during this behavior with high accuracy. The birds were found to fly at much higher speeds during 'screaming parties' than during migration or roosting, on average twice as fast, 20.9  ms⁻¹ (±5.1  ms⁻¹) in horizontal speed. The highest record was 31.1  ms⁻¹ which is the highest measured yet for a swift in self powered flight. Furthermore, the birds were performing steep climbing flights, on average 4.0  ms⁻¹ (±2.8  ms⁻¹) in vertical velocity. A clear trade-off between horizontal speed and vertical speed was found, suggesting that the birds are operating at their maximum.     

The evolution of high summit metabolism and cold tolerance in birds and its impact on present-day distributions

Summit metabolic rate ( M_sum , maximum cold-induced metabolic rate) is positively correlated with cold tolerance in birds, suggesting that high M_sum is important for residency in cold climates. However, the phylogenetic distribution of high M_sum among birds and the impact of its evolution on current distributions are not well understood. Two potential adaptive hypotheses might explain the phylogenetic distribution of high M_sum among birds. The cold adaptation hypothesis contends that species wintering in cold climates should have higher M_sum than species wintering in warmer climates. The flight adaptation hypothesis suggests that volant birds might be capable of generating high M_sum as a byproduct of their muscular capacity for flight; thus, variation in M_sum should be associated with capacity for sustained flight, one indicator of which is migration. We collected M_sum data from the literature for 44 bird species and conducted both conventional and phylogenetically informed statistical analyses to examine the predictors of M_sum variation. Significant phylogenetic signal was present for log body mass, log mass-adjusted M_sum , and average temperature in the winter range. In multiple regression models, log body mass, winter temperature, and clade were significant predictors of log M_sum . These results are consistent with a role for climate in determining M_sum in birds, but also indicate that phylogenetic signal remains even after accounting for associations indicative of adaptation to winter temperature. Migratory strategy was never a significant predictor of log M_sum in multiple regressions, a result that is not consistent with the flight adaptation hypothesis.     

Flight speeds and climb rates of Brent Geese: mass-dependent differences between spring and autumn migration

Aerodynamic theories of bird flight predict that horizontal flight speed will increase with increasing load whereas vertical flight speed will decrease. Horizontal flight speed for birds minimizing overall time on migration is predicted to be higher than flight speed for birds minimizing energy expenditure. In this study we compare flight speeds of Brent Geese Branta b. bernicla recorded by tracking radar and optical range finder during spring and autumn migration in southernmost Sweden, testing the above-mentioned predictions. Geese passing Sweden in spring are substantially heavier than in autumn and there might also be a stronger element of time-selection in spring than in autumn. Recorded airspeeds were significantly higher in spring (mean 19.0 m s⁻¹) than in autumn (mean 17.3 m s⁻¹), the average difference being slightly larger than predicted due to the mass difference alone. The effects on airspeed of wind, vertical speed, flock size and altitude were also analysed, but none of these factors could explain the seasonal difference in airspeed. Hence, the results support the hypothesis of mass-dependent flight speed adjustment. The difference between the two seasons was not large enough to corroborate the hypothesis of a stronger element of time-selection in spring, but this hypothesis cannot be rejected. Vertical flight speeds were lower in spring than in autumn, supporting a negative effect of load on birds' flight power margin.     

Field metabolic rates of black-browed albatrosses Thalassarche melanophrys during the incubation stage

Field metabolic rates (FMR) and activity patterns of black-browed albatrosses Thalassarche melanophrys were measured while at sea and on nest during the incubation stage at Kerguelen Island, southwestern Indian Ocean. Activity-specific metabolic rates of five albatrosses at sea (FMR_at-sea) were measured using doubly labeled water (DLW), and by equipping birds with wet-dry activity data loggers that determined when birds were in flight or on the water. The metabolic rates of four birds incubating their eggs (FMR_on-nest) were also measured using DLW. The mean±SD FMR_at-sea of albatrosses was 611±96 kJ kg⁻¹ d⁻¹ compared to FMR_on-nest of 196±52 kJ kg⁻¹ d⁻¹. While at sea, albatrosses spent 52.9±8.2% (N=3) of their time in flight and they landed on the water 41.2±13.9 times per day. The FMR of black-browed albatrosses appear to be intermediate to that of three other albatross species. Based on at-sea activity, the power requirement of flight was estimated to be 8.7 W kg⁻¹ (or 4.0×predicted BMR), which is high compared to other albatross species, but may be explained by the high activity levels of the birds when at sea. The FMR_at-sea of albatrosses, when scaled with body mass, are lower than other seabirds of similar body size, which probably reflects the economical nature of their soaring flight.     

Gait transition and oxygen consumption in swimming striped surfperch Embiotoca lateralis Agassiz

A flow-through respirometer and swim tunnel was used to estimate the gait transition speed ( U _p-c) of striped surfperch Embiotoca lateralis , a labriform swimmer, and to investigate metabolic costs associated with gait transition. The U _p-c was defined as the lowest speed at which fish decrease the use of pectoral fins significantly. While the tail was first recruited for manoeuvring at relatively low swimming speeds, the use of the tail at these low speeds [as low as 0·75 body (fork) lengths s⁻¹, L _F s⁻¹) was rare (<10% of the total time). Tail movements at these low speeds appeared to be associated with occasional slow manoeuvres rather than providing power. As speed was increased beyond U _p-c, pectoral fin (PF) frequencies kept increasing when the tail was not used, while they did not when PF locomotion was aided by the tail. At these high speeds, the tail was employed for 40–50% of the time, either in addition to pectoral fins or during burst-and-coast mode. Oxygen consumption increased exponentially with swimming speeds up to gait transition, and then levelled off. Similarly, cost of transport ( C _T) decreased with increasing speed, and then levelled off near U _p-c. When speeds ≥ U _p-c are considered, C _T is higher than the theoretical curve extrapolated for PF swimming, suggesting that PF swimming appears to be higher energetically less costly than undulatory swimming using the tail.     

The energetics of mimicry: the cost of pedestrian transport in a formicine ant and its mimic, a clubionid spider

ABSTRACT. Little information exists on the energetics of locomotion in small insects, and none in small spiders. We examined standard rate of oxygen consumption (SO₂) and net cost of transport (NCOT) in Camponotus sericeiventris (Guerin), a formicine ant, Myrmecotypus rettenmeyeri (Unzicker), its clubionid spider mimic (mean masses 43 and 24 mg, respectively), and an unrelated clubionid, Clubiona barroana (Chickering), mean mass 37mg. All species are from Barro Colorado Island, Panama. NCOT in the species studied was 12, 21 and 27 ml O₂g^-1 km^-1, respectively; NCOT in the two spiders was significantly higher than in the ant. The minimum cost of transport of M. rettenmeyeri (8.6ml O₂g^-1 km^-1), though low, did not differ significantly from the value predicted for an insect of its body mass, while its Y intercept elevation (estimated O₂ at zero running speed/SO₂) was extremely high. Y intercept elevation may be high in spiders as a group, leading to high NCOT at low running speeds, and hence to a tendency to adopt ambush predation. It is also predicted that, to minimize locomotion cost per unit distance, spiders will travel faster than insects of equivalent body mass.     

Flight speeds among bird species: allometric and phylogenetic effects

Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)^1/6 and (wing loading)^1/2 among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, U_e) of 138 species, ranging 0.01–10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of U_e in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in U_e. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in U_e. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading.     

Morphology, Velocity, and Intermittent Flight in Birds

Body size, pectoralis composition, aspect ratio of the wing,and forward speed affect the use of intermittent flight in birds.During intermittent non-flapping phases, birds extend theirwings and glide or flex their wings and bound. The pectoralismuscle is active during glides but not during bounds; activityin other primary flight muscles is variable. Mechanical power,altitude, and velocity vary among wingbeats in flapping phases;associated with this variation are changes in neuromuscularrecruitment, wingbeat frequency, amplitude, and gait. Speciesof intermediate body mass (35–158 g) tend to flap-glideat slower speeds and flap-bound at faster speeds, regardlessof the aspect ratio of their wings. Such behavior may reducemechanical power output relative to continuous flapping. Smallerspecies (<20 g) with wings of low aspect ratio may flap-boundat all speeds, yet existing models do not predict an aerodynamicadvantage for the flight style at slow speeds. The behaviorof these species appears to be due to wing shape rather thanpectoralis physiology. As body size increases among species,percent time spent flapping increases, and birds much largerthan 300 g do not flap-bound. This pattern may be explainedby adverse scaling of mass-specific power or lift per unit poweroutput available from flight muscles. The size limit for theability to bound intermittently may be offset somewhat by thescaling of pectoralis composition. The percentage of time spentflapping during intermittent flight also varies according toflight speed.     

Flight performance, echolocation and foraging behaviour in noctule bats Nyctalus noctula

The noctule Nyctulus noctula (Schreber, 1774) is a relatively large (c. 25 g) insectivorous bat which catches insects on the wing (by aerial hawking). Emergence at a maternity roost was earliest relative to sunset when females were lactating, and bats may then have risked predation by flying at higher light levels during a period of high energy demand. Flight performance was quantified by using stereophotogrammetry. At feeding sites bats flew at 6.0 ± 2.1 m/s. This was faster than predicted minimum power speed (V _mp), and either between V _mp and maximum range speed (V _mr), or close to their predicted V _mr, depending on which aerodynamic model of flight power requirements was used. The echolocation behaviour of noctules is flexible. Long duration, low frequency calls (c. 20 kHz) with little frequency modulation were emitted while cruising, but at foraging sites the calls became more frequency-modulated. As the noctule is traditionally thought of as using low frequency echolocation, it was expected to receive weak echoes from small targets and therefore to specialize in eating large insect prey. Although the bats ate mainly beetles, large numbers of small dipterans were also eaten. The noctule is probably able to detect such small items because, when foraging, its calls become broadband and sweep from high frequencies. Higher harmonics are also present, and these may assist in the detection of small prey. In noctules, as in many bats, there appears to be a 1:1 link between wingbeat and call production during the search phase of foraging.     

Wing kinematics of avian flight across speeds

To test whether wing shape affects the kinematics of wing motion during bird flight, we recorded high-speed video (250 Hz) of four species flying in a variable-speed wind tunnel. The birds flew at intervals of 2 m s⁻¹, ranging from 1 m s⁻¹ up to their respective maximum flight speed, which varied from 14 to 17 m s⁻¹ depending on the species. Kinematic data obtained from two synchronized, high-speed video cameras were analyzed using 3D reconstruction. Three species with relatively pointed, high-aspect ratio wings changed wingbeat styles according to flight speed (budgerigar, Melopsittacus undulatus ; cockatiel, Nymphicus hollandicus ; ringed turtle dove, Streptopelia risoria ). These species used a wing-tip reversal upstroke, characterized by supination of the distal wing at mid-upstroke, at equivalent airspeeds ≤7 to 9 m s⁻¹. In faster flight, they used a swept-wing upstroke, without distal wing supination. At mid-upstroke at any speed, wingspan in these species was greater than wrist span. In contrast, at all steady flight speeds, the black-billed magpie Pica hudsonia with relatively broad, low-aspect ratio wings, used a flexed-wing, feathered upstroke in which wrist spans were equal to or greater than wingspans. Our results demonstrate that wing kinematics vary gradually as a function of flight speed, and that the patterns of variation are strongly influenced by external wing shape.     

Hunting flight speeds of five southern African raptors

The flight speeds of hunting falconry birds were determined using global positioning system data loggers. Until now, the hunting flight speed of African raptors has not been directly measured. We predicted that hunting flight speeds would differ between species and that flight dynamics, such as altitude, and bird morphology, particularly wing surface area, would influence maximum and mean flight speeds. This study considered five African raptor species, which included two long-wing species, Lanner Falcon Falco biarmicus and Peregrine Falcon F. peregrinus, one short-wing species, Black Sparrowhawk Accipiter melanoleucus, and two broad-wing species, African Hawk-eagle Aquila spilogaster and Jackal Buzzard Buteo rufofuscus. Maximum and mean hunt speeds differed significantly between the long- and short-wing species. There was no difference in acceleration or deceleration rates between these species, but this could be due to small sample sizes. There was a significant positive correlation between maximum hunt speed and maximum flight height for the long-wing species. Maximum and mean flight speeds were significantly negatively correlated with wing area for all five species in this study. However, following phylogenetic correction, no significant relationship between wing area and maximum hunt speeds was found. This study presents baseline data of hunting speeds in African raptors and further highlights the importance of inter-species variation, which can provide accuracy to flight speed models and the understanding of hunting strategies.     

Energy expenditure during level locomotion in large desert ungulates: the one-humped camel and the domestic donkey

This study sought to quantify the rate of energy expenditure (     ), the total cost of transport (COT_tot) and the net cost of transport (COT_net) in camels Camelus dromedaries and donkeys Equus asinus during level locomotion.     of camels and domestic donkeys were measured at exercise speeds between 0 and 4.17 m s⁻¹. Resting     for camels was significantly ( P <0.05) lower than predicted, while donkeys exhibited resting values similar to mammals of the same body mass. In both camels and donkeys     increased in a nearly linear fashion over the range of exercise speeds. The minimum COT_tot of camels in the walking and pacing gaits were not significantly different ( P =0.27). Similarly, donkeys exhibited no significant difference ( P =0.09) in the minimum COT_tot while walking and trotting. In both camels and donkeys, the minimum COT_tot was significantly ( P <0.05) lower than the predicted COT_tot for mammals of the same body mass. The COT_net in both camels and donkeys was determined to be gait dependent and significantly ( P <0.05) lower than the predicted minimum COT_net values for walking and running. The low COT seen in camels and donkeys results in energy and water savings.     

Gait transition speed, pectoral fin-beat frequency and amplitude in Cymatogaster aggregata, Embiotoca lateralis and Damalichthys vacca

Surfperches are labriform swimmers and swim primarily with their pectoral fins, using the tail to assist only at higher speeds. The transition, from pectoral to pectoral and caudal fins, occurs at a threshold speed that has been termed physiologically and biomechanically 'equivalent' for fishes of different size. The gait transition ( U _P-C) of Cymatogaster aggregata occurred at a higher speed (measured in bodylengths s⁻¹) for smaller fish than larger fish. At U _P-C, pectoral fin-beat frequency was size-dependent: smaller fish have a higher pectoral fin-beat frequency than larger fish. In contrast, at low speeds (i.e. <60% of U _P-C) the pectoral fin-beat frequency was independent of the size of the fish. Inter-specific comparisons of U _P-C, pectoral fin-beat frequency and amplitude among C. aggregata, Embiotoca lateralis and Damalichthys vacca showed that C. aggregata had a higher U _P-C than E. lateralis and D. vacca . The pectoral fin-beat frequency at U _P-C showed no significant differences among species. Cymatogaster aggregata achieved higher U _P-C, in part, through increased fin beat amplitude rather than frequency. These differences in performance may be related to the different habitats in which these species live.     

Kinematics of swimming of penguins at the Detroit Zoo

Kinematic parameters were examined in a study of the swimming abilities of seven species of penguins housed at the Detroit Zoo. Penguins produce thrust over both halves of the wing stroke cycle, as observed in fishes using the caudal or pectoral fins for locomotion, but not in other birds in level forward flight. Unpowered gliding phases between wing strokes were observed in all species at swimming speeds less than 1.25 m/sec, while Emperor, King and Adelie penguins interpose gliding phases over a broad range of speeds. Videotape records reveal that length-specific speed is correlated with increases in wingbeat frequency and, for most of the species examined, stride length. These findings are in contrast to those reported for other, flying birds, which maintain a relatively constant wingbeat frequency but vary stride length with forward speed, and for most fishes, which vary speed with tailbeat frequency but maintain a constant stride length. The results are somewhat comparable to those reported for Cymatogaster , a fish which uses the pectoral fins for locomotion. Drag coefficients of three gliding Emperor penguins were 2.1, 3.0 and 3.0 × 10-⁻³ at Reynolds numbers of 1.25, 1.62 and 1.76 × 10⁶, respectively.     

Endurance at intermediate swimming speeds of Atlantic mackerel, Scomber scombrus L., herring, Clupea harengus L., and saithe, Pollachius virens L.

Endurance and swimming speed were measured in mackerel, herring and saithe when they were induced by the optomotor response to swim at prolonged speeds along a 28-m circular track through still water in a 10-m diameter gantry tank. The maximum sustained swimming speed ( U _ms was measured as body lengths per second ( b.l.s ⁻¹) for each species and for saithe of different size groups. Herring with U _ms of 4.06 b.l.s ⁻¹ (25.3 cm, 13.5°C) were the fastest, mackerel U _ms was 3.5 b.l.s ¹ (33 cm, 11.7°C) and saithe (14.4°C) showed a size effect where U _ms at 25 cm was 3.5 b.l.s ¹ and at 50 cm 2.2 b.l.s ¹. When swimming at speeds higher that U _ms, all three species showed reduced endurance as speed increased. How the curved track reduces the swimming speed is discussed.     

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.     

The effect of wind speed on the flight responses of tsetse flies to CO2: a wind-tunnel study

Abstract. Female Glossina morsitans morsitans Westwood were video-recorded in a wind-tunnel as they entered, in cross-wind flight, a broad plume of CO₂ (a component of host odour). At a wind speed that corresponds with peak catches in the field (c. 0.6 ms-¹) odour produced both significant upwind turning responses (in-flight anemotaxis) and kinetic responses (reduced flight speed and increased sinuosity (m-¹). At a wind speed of c. 0.2 ms-¹ flies displayed anemotactic, but not kinetic, responses to odour. At very low wind speeds (0.1ms-¹) neither upwind turning responses nor kinetic responses to odour were detected. The results are discussed with regard to current theory of host-location by tsetse.     

Pedestrian locomotion energetics and gait characteristics of a diving bird, the great cormorant, Phalacrocorax carbo

Great cormorants Phalacrocorax carbo are foot propelled diving birds that seem poorly suited to locomotion on land. They have relatively short legs, which are presumably adapted for the generation of high forces during the power stroke of aquatic locomotion, and walk with a pronounced "clumsy waddle". We hypothesise (1) that the speed, independent minimum cost of locomotion (C min, ml O2 m(-1)) will be high for cormorants during treadmill exercise, and (2) that cormorants will have a relatively limited speed range in comparison to more cursorial birds. We measured the rate of oxygen consumption (V02) of cormorants during pedestrian locomotion on a treadmill, and filmed them to determine duty factor (the fraction of stride period that the foot is in contact with the ground), foot contact time (tc), stride frequency (f), swing phase duration and stride length. C min was 2.1-fold higher than that predicted by their body mass and phylogenetic position, but was not significantly different from the C min of runners (Galliformes and Struthioniformes). The extrapolated gamma-intercept of the relationship between V02 and speed was 1.9-fold higher than that predicted by allometry. Again, cormorants were not significantly different from runners. Contrary to our hypothesis, we therefore conclude that cormorants do not have high pedestrian transport costs. Cormorants were observed to use a grounded gait with two double support phases at all speeds measured, and showed an apparent gait transition between 0.17 and 0.25 m s(-1). This transition occurs at a Froude number between 0.016 and 0.037, which is lower than the value of approximately 0.5 observed for many other species. However, despite the use of a limited speed range, and a gait transition at relatively low speed, we conclude that the pedestrian locomotion of these foot propelled diving birds is otherwise generally similar to that of cursorial birds at comparable relative velocities.     

Flight speed of seabirds in relation to wind speed and direction

We studied flight speed among all major seabird taxa. Our objectives were to provide further insight into dynamics of seabird flight and to develop allometric equations relating ground speed to wind speed and direction for use in adjusting seabird density estimates (calculated from surveys at sea) for the effect of bird movement. We used triangulation at sea to estimate ground speeds of 1562 individuals of 98 species. Species sorted into 25 “groups” based on similarity in ground speeds and taxonomy. After they were controlled for differences inground speed, the 25 groups sorted into eight major “types” on the basis of response to wind speed and wind direction. Wind speed and direction explained 1664% of the variation in ground speed among seabird types. For analyses on air speed (ground speed minus apparent wind speed), we divided the 25 groups according to four flight styles: gliding, flap-gliding, glide-flapping and flapping. Tailwind speed had little effect on air speed of gliders (albatrosses and large gadfly petrels), but species that more often used flapping decreased air speed with increase in tailwinds. All species increased air speeds significantly with increased headwinds. Gliders showed the greatest increase relative to increase in headwind speed and flappers the least. With tailwind flight, air speeds were greatest among species with highest wing loading for each flight style except gliders, which showed no relationship. For headwind flight, species with higher wing loading had higher air speeds; however, the relation was weaker in flappers compared with species using some amount of gliding. In contrast, analyses for air speed ratio (i.e. difference between air speed in acrosswinds [with no apparent wind] and speed flown into headwinds, or with tailwinds, divided by speed acrosswind) revealed that among species using some flapping, and with lower wing loading (surface-feeding shearwaters, small gadfly petrels, storm petrels, phalaropes, gulls and terns), adjusted air speeds more than those with higher wing loading (alcids, “diving shearwaters”, “Manx-type shearwaters”, pelicans, boobies and cormorants). As a result, most flappers of low wing loading flew much faster than V_mr (the most energy efficient air speed per distance flown) when flying into headwinds. We suggest that better-than-predicted gliding performance with acrosswinds and tailwinds of large gadfly petrels, compared with albatrosses, resulted from a different type of “soaring” not previously described in seabirds.