The physiology and biomechanics of avian flight at high altitude
Abstract
Many birds fly at high altitude, either during long-distance flights or by virtue of residence in high-elevation habitats. Among the many environmental features that vary systematically with altitude, five have significant consequences for avian flight performance: ambient wind speeds, air temperature, humidity, oxygen availability, and air density. During migratory flights, birds select flight altitudes that minimize energy expenditure via selection of advantageous tail- and cross-winds. Oxygen partial pressure decreases substantially to as little as 26% of sea-level values for the highest altitudes at which birds migrate, whereas many taxa reside above 3000 meters in hypoxic air. Birds exhibit numerous adaptations in pulmonary, cardiovascular, and muscular systems to alleviate such hypoxia. The systematic decrease in air density with altitude can lead to a benefit for forward flight through reduced drag but imposes an increased aerodynamic demand for hovering by degrading lift production and simultaneously elevating the induced power requirements of flight. This effect has been well-studied in the hovering flight of hummingbirds, which occur throughout high-elevation habitats in the western hemisphere. Phylogenetically controlled studies have shown that hummingbirds compensate morphologically for such hypodense air through relative increases in wing size, and kinematically via increased stroke amplitude during the wingbeat. Such compensatory mechanisms result in fairly constant power requirements for hovering at different elevations, but decrease the margin of excess power available for other flight behaviors.
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Claims the paper makes
1 of 94 cited sources checked · 1 supported · 0 flagged
- Premisecitation
Research on avian flight has typically been conducted at or near sea-level conditions for anthropogenic reasons deriving from contemporary altitudinal distribution of modern humans.
“Research on avian flight has, for anthropogenic reasons deriving from contemporary altitudinal distribution of modern humans (Cohen and Small, 1998), typically been conducted at or near sea-level conditions.”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Cohen and Small, 1998 source
- Findingown data
Stroke amplitude during hovering exhibited a systematic increase among species at higher elevations.
“Stroke amplitude during hovering exhibited a systematic increase among species at higher elevations”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Altshuler and Dudley, 2003 source
- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingreasoning
The power reserve systematically declines at higher elevation.
“The power reserve thus systematically declines at higher elevation (Fig. 2)”
- Also stated in: § Conclusions · plain
“Hummingbird power margin decreases with elevation.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Also stated in: § Conclusions · plain
- Findingown data
Hummingbirds failed to perform sustained hovering at specific oxygen- or density-equivalent elevations during density- and oxygen-reduction trials.
“Oxygen-or density-equivalent elevations at which hummingbirds failed to perform sustained hovering during density-and oxygen-reduction trials*”
Quote in the paperSubstantial claim - Findingcitation
Load-lifting elicits maximum stroke amplitudes in all taxa.
“load-lifting elicits maximum stroke amplitudes in all taxa”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingcitation
Body size systematically increases among hummingbird species at higher elevations.
“Body size systematically increases among hummingbird species at higher elevations (Altshuler et al., 2004b; Fig. 1)”
- Also stated in: § Conclusions · plain
“Hummingbird body mass increases with elevation.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Also stated in: § Conclusions · plain
- Premisecitation
High-elevation taxa such as the bar-headed goose possess hemoglobin with higher baseline affinity for oxygen.
“but high-elevation taxa such as the bar-headed goose possess hemoglobin with higher baseline affinity for oxygen (Black and Tenney, 1980).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Black and Tenney, 1980 source
- Interpretationreasoning
Such compensatory mechanisms result in fairly constant power requirements for hovering at different elevations.
“Such compensatory mechanisms result in fairly constant power requirements for hovering at different elevations, but decrease the margin of excess power available for other flight behaviors.”
Quote in the paperSubstantial claim - Premisenone given
Air density systematically declines with elevation in proportion to the concomitant change in total pressure.
“As with oxygen partial pressure, air density systematically declines with elevation in proportion to the concomitant change in total pressure.”
Quote in the paperNot a substantial claim - Premisereasoning
Gas exchange between the avian pulmonary and circulatory systems is particularly efficient due to a suite of anatomical adaptations distinguishing avian gas exchange from mammalian counterparts.
“Gas exchange between the avian pulmonary and circulatory systems is particularly efficient due to a suite of anatomical adaptations distinguishing avian gas exchange from mammalian counterparts.”
Quote in the paperSubstantial claim - Premisenone given
Oxygen partial pressure decreases substantially to as little as 26% of sea-level values for the highest altitudes at which birds migrate.
“Oxygen partial pressure decreases substantially to as little as 26% of sea-level values for the highest altitudes at which birds migrate, whereas many taxa reside above 3000 meters in hypoxic air.”
Quote in the paperNot a substantial claim - Premisereasoning
The systematic decrease in air density with altitude can lead to a benefit for forward flight through reduced drag.
“The systematic decrease in air density with altitude can lead to a benefit for forward flight through reduced drag but imposes an increased aerodynamic demand for hovering by degrading lift production and simultaneously elevating the induced power requirements of flight.”
Quote in the paperSubstantial claim - Premisecitation
Muscle capillary-per-fiber number is higher for birds at high elevations in both highly aerobic pectoral muscles and less aerobic leg muscles.
“Specifically, muscle capillary-per-fiber number is higher for birds at high elevations in both highly aerobic pectoral muscles and less aerobic leg muscles (Hepple et al., 1998;Mathieu-Costello et al., 1998).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Premisereasoning
Hummingbirds compensate kinematically for hypodense air via increased stroke amplitude during the wingbeat.
“and kinematically via increased stroke amplitude during the wingbeat.”
Quote in the paperSubstantial claim - Interpretationreasoning
The high aerobic demands of flight lie at odds with reduced oxygen availability at high altitude.
“The high aerobic demands of flight thus lie at odds with reduced oxygen availability at high altitude.”
Quote in the paperSubstantial claim - Premisereasoning
The systematic decrease in air density with altitude imposes an increased aerodynamic demand for hovering by degrading lift production and elevating induced power requirements.
“but imposes an increased aerodynamic demand for hovering by degrading lift production and simultaneously elevating the induced power requirements of flight.”
Quote in the paperSubstantial claim - Premisecitation
Phylogenetically controlled studies have shown that hummingbirds compensate morphologically for hypodense air through relative increases in wing size.
“Phylogenetically controlled studies have shown that hummingbirds compensate morphologically for such hypodense air through relative increases in wing size, and kinematically via increased stroke amplitude during the wingbeat.”
Quote in the paperSubstantial claim - Premisecitation
During deep hypoxia, respiration is ultimately limited by blood perfusion.
“During deep hypoxia, however, respiration is ultimately limited by blood perfusion (Fedde et al., 1989;Shams and Scheid, 1989).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Premisenone given
Ambient wind speed generally increases with altitude.
“A second feature of aerodynamic significance is the general trend of increasing ambient wind speed with altitude.”
Quote in the paperNot a substantial claim - Interpretationreasoning
Such compensatory mechanisms decrease the margin of excess power available for other flight behaviors.
“but decrease the margin of excess power available for other flight behaviors.”
Quote in the paperSubstantial claim - Premisecitation
Arterial hypocapnia does not result in vasoconstriction in birds.
“Whereas this condition leads to arterial constriction and decreased blood flow in many mammals, arterial hypocapnia does not result in vasoconstriction in birds (Faraci, 1991), and both low-and high-elevation taxa exhibit normally cognitive function at elevations well above terrestrial habitation (Black and Tenney, 1980).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Faraci, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
The cost of supporting body weight (induced power requirements) increases at lower air density.
“In particular, the cost of supporting body weight (i.e., the induced power requirements) increase at lower air density, whereas profile drag (on the wings) and parasite drag (on the body), together with their associated power expenditures, will concomitantly decrease (Pennycuick, 1975;Norberg, 1990;Rayner, 1990).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Pennycuick, 1975 source
- cites Norberg, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Rayner, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Profile drag on the wings and parasite drag on the body, together with their associated power expenditures, decrease at lower air density.
“whereas profile drag (on the wings) and parasite drag (on the body), together with their associated power expenditures, will concomitantly decrease (Pennycuick, 1975;Norberg, 1990;Rayner, 1990).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Pennycuick, 1975 source
- cites Norberg, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Rayner, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
The concomitant reduction in the Reynolds number of the wings may reduce their effective lift:drag ratio.
“the concomitant reduction in the Reynolds number of the wings may also reduce their effective lift:drag ratio”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Ellington, 1984a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Vogel, 1988
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dudley, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation
The precise magnitude of the effect of lower air density on hovering aerodynamics varies with the particular aerodynamic mechanisms and wing morphologies under consideration.
“The precise magnitude of this effect, however, will vary with the particular aerodynamic mechanisms and wing morphologies under consideration”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Usherwood and Ellington, 2002 source
- cites Altshuler et al., 2004a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
The distributional limit of hummingbirds over 5000 m is most likely determined by ecological rather than physiological features.
“which is a distributional limit most likely determined by ecological rather than physiological features.”
Quote in the paperNot a substantial claim - Premisereasoning
Water loss via respiration increases with altitude due to systematic decrease in relative humidity.
“Water loss via respiration increases with altitude due to systematic decrease in relative humidity.”
Quote in the paperSubstantial claim - Premisenone given
There are few data supporting hypotheses that thermoregulation and water balance are regulated through altitude selection.
“Despite such constraints on thermoregulation and water balance, there are few data supporting hypotheses that these physiological processes are regulated through altitude selection.”
Quote in the paperNot a substantial claim - Premisecitation
Hummingbird flight requires the highest mass-specific oxygen consumption of any vertebrate locomotor mode.
“Hummingbird flight requires the highest massspecific oxygen consumption of any vertebrate locomotor mode (Suarez, 1992), but these birds nevertheless inhabit high elevations throughout the New World (Schuchmann, 1999), with some taxa residing up to 5000 m (Carpenter, 1976;Rahbek and Graves, 2000).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Suarez, 1992 source
- Premisecitation
Lower air density yields reduced forces on wings if kinematics are unchanged.
“Lower air density yields reduced forces on wings if kinematics are unchanged”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Ellington, 1984a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Vogel, 1988
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dudley, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
By compiling several sources of data, Butler et al. (1997) calculated that the large body masses measured at stopover sites were only possible by accounting for wind-assisted flight.
“By compiling several sources of data, Butler et al. (1997) calculated that the large body masses measured at stopover sites were only possible by accounting for wind-assisted flight.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Butler et al., 1997 source
- Interpretationreasoning
The systematic decline in air density with increased elevation adversely affects the aerodynamics of hovering flight.
“The systematic decline in air density with increased elevation must adversely affect the aerodynamics of hovering flight.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Ellington, 1984a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Vogel, 1988
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dudley, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Rests on an assumption: Air density declines systematically with increased elevation.
- Premisereasoning
Currently, there is equivocal evidence that birds actually dehydrate during migratory flights.
“Currently, there is equivocal evidence that birds actually dehydrate during migratory flights.”
Quote in the paperNot a substantial claim - Premisecitation
Intra- and interspecific comparisons of bird taxa suggest relatively larger wings at higher altitudes.
“Intra-and interspecific comparisons of bird taxa suggest relatively larger wings at higher altitudes”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation
Hummingbirds failed in hovering flight at air densities well above those characteristic of failure in normoxic hypobaria when normal air was replaced with pure helium, illustrating a constraint of oxygen delivery.
“Chai and Dudley (1996) replaced normal air with pure helium, thus reducing air density as well as the partial pressure of oxygen.Hummingbirds failed in hovering flight at air densities well above those characteristic of failure in normoxic hypobaria, clearly illustrating a constraint of oxygen delivery.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Chai and Dudley, 1996
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Swift flight altitudes are selected according to temperature, with swifts flying at higher altitudes on warmer nights.
“Instead, flight altitudes are selected according to temperature, with swifts flying and Dudley (1995) and experiments with Selasphorus platycercus and S. rufus were performed by Altshuler and Dudley (2003).”
- Also stated in: § Thermoregulation and water balance · plain
“at higher altitudes on warmer nights (Backman and Alerstam, 2001).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Backman and Alerstam, 2001 source
- Also stated in: § Thermoregulation and water balance · plain
- Premisecitation
Relatively larger wings at higher altitudes are systematically demonstrated among hummingbird species.
“an effect systematically demonstrated among hummingbird species”
Quote in the paperNot a substantial claim0 of 2 sources checked - Interpretationcitation
Relative wing size will increase at higher altitudes to offset the increased induced power requirements associated with lower air densities.
“One immediate biomechanical prediction is that relative wing size will increase at higher altitudes to offset the increased induced power requirements associated with lower air densities”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ellington, 1984b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Along an altitudinal cline in the Canadian Arctic, migrating birds' flight speed increased by the predicted amount after accounting for the effects of wind speed and direction.
“Along this altitudinal cline, flight speed increased by the predicted amount after accounting for the effects of wind speed and direction.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Hedenström et al., 2002 source
- Premisecitation
At high altitude, energetic costs may increase substantially depending on the relative magnitude of the induced and profile power components of total mechanical power expenditure.
“Also at high altitude, energetic costs may increase substantially depending on the relative magnitude of the induced and profile power components of total mechanical power expenditure”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Ellington, 1984a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Norberg, 1990
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation
Benefits derived from wind assistance are likely to motivate migrants to a greater extent than thermoregulation or water balance.
“Instead, benefits derived from wind assistance are likely to motivate migrants to a greater extent (Liechti et al., 2000), although this depends on the importance attributed to maintaining water balance (Klaassen and Biebach, 2000).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Liechti et al., 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Klaassen and Biebach, 2000 source
- Premisecitation
Some migrants would ascend up to 9000 m to encounter air jets in which they could fly with groundspeeds greater than 45 meters/second.
“Furthermore, some migrants would ascend up to 9000 m to encounter air jets in which they could fly with groundspeeds greater than 45 meters/second (Liechti and Schaller, 1999).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Liechti and Schaller, 1999 source
- Premisecitation
Of all meteorological variables, only tailwind velocity was significantly correlated with the altitude of migratory flights.
“Of all meteorological variables, Bruderer et al. (1995) found that only tailwind velocity was significantly correlated with the altitude of migratory flights.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bruderer et al., 1995 source
- Interpretationcitation
The systematic decline in hummingbird power reserves with elevation suggests that the physically imposed consequences of high-elevation residence impinge on a diversity of flight behaviors.
“The systematic decline in hummingbird power reserves with elevation (Altshuler et al., 2004b, see below) suggests that the physically imposed consequences of high-elevation residence impinge on a diversity of flight behaviors.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premiseown data
In all hypoxic and hypodense air experiments to date, hummingbirds failed to hover at equivalent elevations over 5000 m.
“However, it is important to note that in all hypoxic and hypodense air experiments to date, hummingbirds failed to hover at equivalent elevations over 5000 m (Table 1), which is a distributional limit most likely determined by ecological rather than physiological features.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Table 1
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
This systematic change in nectar availability with altitude could well alter the foraging ecology of hummingbirds, given increases in thermoregulatory demand and costs of hovering flight at higher elevations.
“This systematic change in nectar availability with altitude could well alter the foraging ecology of hummingbirds, given increases in thermoregulatory demand and costs of hovering flight at higher elevations.”
Quote in the paperSubstantial claim - Premisecitation
The 20-26 g Giant Hummingbird (Patagona gigas) is resident at elevations up to 4000 m.
“the 20-26 g Giant Hummingbird (Patagona gigas) is resident at elevations up to 4000 m”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ortiz-Crespo, 1974 source
- Interpretationreasoning
Low-density air at high altitudes will affect forward flight and hovering differently.
“Low-density air at high altitudes will affect forward flight and hovering differently.”
Quote in the paperSubstantial claim - Premisecitation
Manipulations of hovering flight in hummingbirds using hypodense helium mixtures under both normoxic and hypoxic conditions revealed that low-density air elicits a systematic increase in stroke amplitude but essentially unchanged wingbeat frequencies.
“Manipulations of hovering flight in hummingbirds using hypodense helium mixtures under both normoxic and hypoxic conditions (Chai andDudley, 1995, 1996;Altshuler and Dudley, 2003) similarly revealed that low-density air elicits a systematic increase in stroke amplitude but essentially unchanged wingbeat frequencies.”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationreasoning
For forward flight, hypodense air reduces drag forces on the wings and body, which can result in faster flight speeds.
“For the former, hypodense air reduces drag forces on the wings and body, which can result in faster flight speeds.”
Quote in the paperSubstantial claim - Premisecitation
The adverse effects of air density on lift and power production will be systematically more pronounced at greater body mass.
“the adverse effects of air density on lift and power production will be systematically more pronounced at greater body mass”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Norberg, 1995 source
- Premisenone given
Studies of flight aerodynamics in hypodense air have been, with the exception of Tucker (1968b), limited to hovering hummingbirds.
“Studies of flight aerodynamics in hypodense air have been, with the exception of the aforementioned work by Tucker (1968b), limited to hovering hummingbirds.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Tucker, 1968b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Physical constraints imposed by low-density high-elevation air are most evident aerodynamically during hovering flight, during which lift and power requirements of the flight motor are extreme.
“Physical constraints imposed by low-density highelevation air are most evident aerodynamically during hovering flight, during which lift and power requirements of the flight motor are extreme”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ellington, 1984a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Lower nectar temperatures will impose additional energetic costs on meal acquisition by hummingbirds.
“lower nectar temperatures will impose additional energetic costs on meal acquisition by hummingbirds (Lotz et al., 2003)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Lotz et al., 2003 source
- Premisecitation
A similar response is seen for hovering in hyperoxic but hypodense gas mixtures.
“A similar response is seen for hovering in hyperoxic but hypodense gas mixtures (Chai et al., 1996;Altshuler et al., 2001).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Chai et al., 1996
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Altshuler et al., 2001 source
- Premisecitation
Trochilid species diversity, with over 320 taxa, is greatest over the altitudinal range of 1500-2500 m.
“Trochilid species diversity, with over 320 taxa, is greatest over the altitudinal range of 1500-2500 m”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Schuchmann, 1999
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dudley, 2001 source
- Premisecitation
The relative mechanical and metabolic capacities of flying animals tend to decline with increasing size.
“the relative mechanical and metabolic capacities of flying animals tend to decline with increasing size”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation
Sucrose concentrations of hummingbird-pollinated flowers decrease across elevations for sites in the southwestern USA and in Costa Rica.
“sucrose concentrations of hummingbird-pollinated flowers decreases across elevations for sites in the southwestern USA (Pyke and Waser, 1981;Cruden et al., 1983) and in Costa Rica (Hainsworth and Wolf, 1972;Baker, 1975;Pyke and Waser, 1981)”
Quote in the paperNot a substantial claim0 of 3 sources checked - Interpretationreasoning
The presence of larger hummingbird species at high altitudes exacerbates the demands of hovering flight.
“Some of the larger species are, somewhat paradoxically, most common at high altitudes, exacerbating the demands of hovering flight.”
Quote in the paperSubstantial claim- Rests on an assumption: Some of the larger hummingbird species are most common at high altitudes.
- Interpretationown data
Kinematic responses by individual birds to hypodense air are mirrored on evolutionary timescales among species.
“indicating that kinematic responses by individual birds to hypodense air are mirrored on evolutionary timescales among species”
Quote in the paperSubstantial claim - Interpretationreasoning
High-elevation hummingbirds are not limited in their capacity for normal hovering flight despite the challenges imposed by hypobaric environments.
“High-elevation hummingbirds are thus not limited in their capacity for normal hovering flight despite the challenges imposed by hypobaric environments.”
Quote in the paperSubstantial claim - Interpretationcitation
The interspecific increase in stroke amplitude of hovering hummingbirds at higher elevations, when coupled with the geometrical constraint on wing motions to amplitudes of approximately 180 on either side of the body, has important implications for aerodynamic and energetic reserves during flight.
“The interspecific increase in stroke amplitude of hovering hummingbirds at higher elevations, when coupled with the geometrical constraint on wing motions to amplitudes of approximately 180 on either side of the body (Chai and Dudley, 1995), has important implications for aerodynamic and energetic reserves during flight.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Chai and Dudley, 1995 source
- Interpretationreasoning
The decline in power reserve possibly compromises performance in varied contexts of both natural and sexual selection.
“possibly compromising performance in varied contexts of both natural and sexual selection”
Quote in the paperSubstantial claim
- May combine two claims. 29
- Classified as probably not a substantial claim. 21
May combine two claims.
FindingcitationAmong species, the mass-specific aerodynamic power requirements for hovering flight are approximately constant with respect to altitude, because of systematic interspecific increases in relative wing size and in stroke amplitude at higher elevations.
“Among species, the mass-specific aerodynamic power requirements for hovering flight are approximately constant with respect to altitude, because of systematic interspecific increases in relative wing size and in stroke amplitude at higher elevations.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
FindingcitationWingbeat frequency declined as predicted on allometric grounds with body mass but was uncorrelated with elevation.
“Wingbeat frequency, by contrast, declined as predicted on allometric grounds with body mass but was uncorrelated with elevation.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Altshuler and Dudley, 2003 source
- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
FindingreasoningHigher-elevation hummingbirds already hover with increased stroke amplitudes, and correspondingly possess less reserve capacity in wing motions and related power production supplemental to that required for normal hovering.
“higher-elevation hummingbirds already hover with increased stroke amplitudes, they correspondingly possess less reserve capacity in wing motions and related power production supplemental to that required for normal hovering”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Altshuler et al., 2004b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Classified as probably not a substantial claim.
Premisenone givenBirds exhibit numerous adaptations in pulmonary, cardiovascular, and muscular systems to alleviate hypoxia.
“Birds exhibit numerous adaptations in pulmonary, cardiovascular, and muscular systems to alleviate such hypoxia.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationMigrating birds may be alternatively impeded or aided by higher wind speeds if the wind direction is against or in the direction of desired forward progress.
“Migrating birds, on the other hand, may be alternatively impeded or aided by higher wind speeds if the wind direction is against or in the direction of desired forward progress, respectively (Green et al., 2004).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Green et al., 2004 source
Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Premisenone givenMany bird taxa are high-elevation specialists, and a large number of migrants fly at substantial altitude.
“Nonetheless, many bird taxa are high-elevation specialists, and a large number of migrants fly at substantial altitude.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Premisenone givenDuring migratory flights, birds select flight altitudes that minimize energy expenditure via selection of advantageous tail- and crosswinds.
“During migratory flights, birds select flight altitudes that minimize energy expenditure via selection of advantageous tail-and crosswinds.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationThe oxygen delivery systems of birds function well across a broad range of oxygen partial pressures and exhibit considerable adaptive plasticity when rapidly exposed to deep hypoxia.
“In summary, the oxygen delivery systems of birds function well across a broad range of oxygen partial pressures and exhibits considerable adaptive plasticity when rapidly exposed to deep hypoxia (Shams and Scheid, 1993).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Shams and Scheid, 1993 source
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Premisenone givenIn contrast with many mammals, birds subjected experimentally to low barometric pressures do not exhibit some of the most maladaptive responses.
“In contrast with many mammals, birds subjected experimentally to low barometric pressures do not exhibit some of the most maladaptive responses.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationBlood leaving the lung interface can have the same oxygen partial pressure as inspired air, indicating that oxygen delivery in birds is not limited by the pulmonary system.
“The net result of these adaptations is that blood leaving the lung interface can have the same oxygen partial pressure as inspired air, indicating that oxygen delivery in birds is not limited by the pulmonary system (Fedde, 1990).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Fedde, 1990 source
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Premisenone givenMany avian taxa reside above 3000 m in hypoxic air.
“whereas many taxa reside above 3000 meters in hypoxic air.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationIn response to hypobaria, birds increase overall rates of oxygen consumption and respiratory rates, whereas these parameters decrease for mice.
“Similarly in response to hypobaria, birds increase overall rates of oxygen consumption and respiratory rates, whereas these parameters decrease for mice (Tucker, 1968b).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tucker, 1968b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
PremisereasoningAerodynamic forces typically vary in linear proportion to air density, so morphological and kinematic compensation is necessary to effect flight at different elevations.
“Because aerodynamic forces typically vary in linear proportion to air density, morphological and kinematic compensation is necessary to effect flight at different elevations.”
Quote in the paperSubstantial claim- Rests on an assumption: Aerodynamic forces vary in linear proportion to air density.
May combine two claims.
Premisenone givenFive environmental features vary systematically with altitude and have significant consequences for avian flight performance: ambient wind speeds, air temperature, humidity, oxygen availability, and air density.
“Among the many environmental features that vary systematically with altitude, five have significant consequences for avian flight performance: ambient wind speeds, air temperature, humidity, oxygen availability, and air density.”
Quote in the paperSubstantial claimMay combine two claims.
Premisenone givenSea-level density of air at 20 C is about 1.21 kg/m3, decreasing to 0.95 kg/m3 at 2000 m elevation and 0.74 kg/m3 at 4000 m, the latter being a 40% reduction relative to sea-level.
“Sea-level density of air at 20 C is about 1.21 kg/m 3 , decreasing to 0.95 kg/m 3 at 2000 m elevation and 0.74 kg/m 3 at 4000 m, the latter being a 40% reduction relative to the sea-level value.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim.
InterpretationreasoningMorphological, behavioral, and physiological adaptations to altitude must correspondingly pertain.
“Morphological, behavioral, and physiological adaptations to altitude must correspondingly pertain.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationAerodynamic models predict an increase in cruising flight speed with altitude at the approximate rate of 5% per 1000 m, primarily because of the reduction in wing and body drag, and because of the relatively small magnitude of induced power in forward flight relative to other components of energetic expenditure.
“Specifically, aerodynamic models predict an increase in cruising flight speed with altitude at the approximate rate of 5% per 1000 m, primarily because of the reduction in wing and body drag, and because of the relatively small magnitude of induced power in forward flight relative to other components of energetic expenditure (Pennycuick, 1978;Hedenstro ¨m, 2003).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationHummingbirds inhabit high elevations throughout the New World, with some taxa residing up to 5000 m.
“but these birds nevertheless inhabit high elevations throughout the New World (Schuchmann, 1999), with some taxa residing up to 5000 m (Carpenter, 1976;Rahbek and Graves, 2000).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Schuchmann, 1999
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Carpenter, 1976
- cites Rahbek and Graves, 2000 source
May combine two claims.
PremisecitationEndurance in the wind tunnel decreased substantially with decreasing pressure, although one bird provided with supplemental oxygen apparently flew better than the birds breathing rarefied air.
“Endurance in the wind tunnel decreased substantially with decreasing pressure, although one bird provided with supplemental oxygen apparently flew better than the birds breathing rarefied air (Tucker, 1968b).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Tucker, 1968b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
PremisecitationIn some taxa, departure probabilities are strongly correlated with the presence of favorable tailwinds, whereas other taxa only depart when winds are absent or weak regardless of wind direction.
“In some cases, departure probabilities are strongly correlated with the presence of favorable tailwinds (e.g., A ˚kesson and Hedenstro ¨m, 2000;Klaassen et al., 2004), whereas other taxa only depart when winds are absent or weak regardless of wind direction (e.g., Schaub et al., 2004).”
Quote in the paperNot a substantial claim0 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenNo one else has used a hypobaric wind tunnel to study forward flight in birds.
“To our knowledge, no one else has used a hypobaric wind tunnel to study forward flight in birds, although this would represent another compelling test of Pennycuick's (1978) hypothesis.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationTheoretical analysis predicts that by accounting for wind conditions, birds could double their ground speed and accordingly save as much as half of the energy required for migratory flights.
“Theoretical analysis also predicts that by accounting for wind conditions, birds could double their ground speed and accordingly save as much as half of the energy required for migratory flights (Liechti and Bruderer, 1998).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Liechti and Bruderer, 1998 source
May combine two claims.
PremisecitationUnder laboratory conditions, rates of oxygen consumption by hovering hummingbirds increase under reduced total pressure and under hypodense but normoxic conditions.
“Under laboratory conditions, rates of oxygen consumption by hovering hummingbirds increases under reduced total pressure (Berger, 1974a,b) and under hypodense but normoxic conditions (Chai and Dudley, 1995).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Berger, 1974a,b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Chai and Dudley, 1995 source
May combine two claims.
PremisecitationProviding supplemental oxygen does not increase aerodynamic performance in hypodense gas, but does allow for longer hovering durations when challenged aerodynamically.
“Providing supplemental oxygen does not increase aerodynamic performance in hypodense gas (Chai et al., 1996), but does allow for longer hovering durations when challenged aerodynamically (Altshuler et al., 2001).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Chai et al., 1996
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Altshuler et al., 2001 source
Classified as probably not a substantial claim.
PremisecitationIn a computer-simulation model, respiratory dehydration would ultimately limit flight duration and distance, particularly in birds with sufficient fat stores to energetically fuel their flight.
“In a computersimulation model, Carmi et al. (1992) concluded that respiratory dehydration would ultimately limit flight duration and distance, particularly in birds with sufficient fat stores to energetically fuel their flight.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Carmi et al., 1992
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenThe effects of hypobaria on more diverse features of the avian flight envelope, including takeoff performance and maneuverability, are unstudied.
“Also unstudied are the effects of hypobaria on more diverse features of the avian flight envelope, including takeoff performance and maneuverability.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim. May combine two claims.
PremisecitationBody water content can remain constant even as both birds and mammals undergo critical water stress, and thus may be a poor predictor of dehydration state.
“However, Klaassen (2004) has pointed out that body water content can remain constant even as both birds and mammals undergo critical water stress, and thus may be a poor predictor of dehydration state.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Klaassen, 2004 source
May combine two claims.
InterpretationreasoningA range of compensatory morphological, physiological, and biomechanical adaptations may have permitted larger hummingbirds to progressively colonize higher elevations in South America, but this outcome has not been entirely without cost.
“A range of compensatory morphological, physiological, and biomechanical adaptations may have permitted larger hummingbirds to progressively colonize higher elevations in South America, but this outcome has not been entirely without cost.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Premisenone givenBiotic factors also vary with elevation, although their influence on flight performance has received less attention than physical factors.
“Biotic factors also vary with elevation, although their influence on flight performance has received less attention than physical factors.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Premisenone givenRespiratory physiology has been the best studied type of such responses, and diverse adaptations are evident in the design of respiratory systems, oxygen-blood transport, and overall hypoxia resistance.
“Respiratory physiology has been the best studied type of such responses, and diverse adaptations are evident in the design of respiratory systems, oxygen-blood transport, and overall hypoxia resistance.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Interpretationnone givenBirds can respond on behavioral, ecological, and evolutionary timescales.
“Birds in turn can respond on behavioral, ecological, and evolutionary timescales.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Premisenone givenAltitudinal gradients in ambient winds are often used by avian migrants to enhance groundspeeds and overall energetic efficiency of migration.
“Altitudinal gradients in ambient winds are often used by avian migrants to enhance groundspeeds and overall energetic efficiency of migration.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based). May combine two claims.
Premisenone givenDuring hovering, drag forces on the body are, by definition, nonexistent and induced power requirements to offset gravity are a dominant avenue of energy expenditure.
“During hovering, drag forces on the body are, by definition, nonexistent and induced power requirements to offset gravity are a dominant avenue of energy expenditure.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationBerger (1974a) experimentally imposed hypobaria on two species of montane hummingbirds, and elicited systematic increases in stroke amplitude and wing angle of attack.
“Berger (1974a) experimentally imposed hypobaria on two species of montane hummingbirds, and elicited systematic increases in stroke amplitude and wing angle of attack”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Berger, 1974a
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationWingbeat kinematics of the Giant Andean Hummingbird are noticeably different at half-stroke transitions relative to those of other hummingbirds.
“Wingbeat kinematics of the Giant Andean Hummingbird are noticeably different at half-stroke transitions relative to those of other hummingbirds”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites M.-J. Fernandez, pers. comm.
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing).
Interpretationnone givenBehavioral means of compensation to the adverse energetics of high-altitude flight are possible.
“Behavioral means of compensation to the adverse energetics of high-altitude flight are also possible.”
Quote in the paperNot a substantial claim- Rests on an assumption: High-altitude hummingbirds often perch while feeding, whereas their lowland counterparts almost never do so.
May combine two claims.
Interpretationnone givenThese effects are likely to be of secondary importance for long-distance avian migrants, but must impinge substantially on the flight biology of high-altitude residents as well as of altitudinal migrants.
“These effects are likely to be of secondary importance for long-distance avian migrants, but must impinge substantially on the flight biology of high-altitude residents as well as of altitudinal migrants.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningHummingbirds (family Trochilidae) represent an ideal taxon with which to evaluate questions of physiological and biomechanical adaptation to high elevation.
“Hummingbirds (family Trochilidae) represent an ideal taxon with which to evaluate such questions of physiological and biomechanical adaptation to high elevation.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based). May combine two claims.
Premisenone givenReductions in oxygen partial pressure, air density, and air temperature all potentially exert major influences on avian flight physiology.
“Reductions in oxygen partial pressure, air density, and air temperature all potentially exert major influences on avian flight physiology.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningReduced air density at high elevation imposes aerodynamic and energetic costs on hovering, but a number of compensatory responses in flight-related morphology and biomechanics have now been demonstrated among different hummingbird species across elevational gradients.
“Accordingly, reduced air density at high elevation imposes aerodynamic and energetic costs on hovering, but a number of compensatory responses in flight-related morphology and biomechanics have now been demonstrated among different hummingbird species across elevational gradients.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
InterpretationreasoningSuch diminished reserves should also be evident in individual birds that migrate seasonally to higher elevations.
“Such diminished reserves should also be evident in individual birds that migrate seasonally to higher elevations.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationA systematic increase in stroke amplitude at constant wingbeat frequency increases relative wing speed to overcome otherwise declining lift production, and simultaneously mitigates the enhanced induced power expenditure associated with hypodense air.
“Overall, such a systematic increase in stroke amplitude at constant wingbeat frequency increase relative wing speed to overcome otherwise declining lift production, and simultaneously mitigate the enhanced induced power expenditure associated with hypodense air”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ellington, 1984b
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Premisenone givenRelevant data are extremely limited, and overall the variation in wind velocity, oxygen partial pressure and in air density with altitude are of primary relevance to the mechanics and physiology of avian flight.
“Relevant data are however extremely limited, and overall the variation in wind velocity, oxygen partial pressure and in air density with altitude are of primary relevance to the mechanics and physiology of avian flight.”
Quote in the paperNot a substantial claim
The paper relies on this without support.
Aerodynamic forces vary in linear proportion to air density.
“Because aerodynamic forces typically vary in linear proportion to air density, morphological and kinematic compensation is necessary to effect flight at different elevations.”
The paper relies on this without support.
The environmental lapse rate in air temperature remains linear within the troposphere.
“The environmental lapse rate in air temperature through adiabatic cooling and increase in water content is about 0.65 C/100 m, and remains linearly so within the troposphere, so that an approximately 26 C difference characterizes air at 4000 m relative to that at sea level.”
The paper relies on this without support.
Laboratory decoupling of air density and oxygen partial pressure can reveal the physiological and aerodynamic consequences of high-altitude flight.
“Because it is possible to experimentally decouple air density and oxygen partial pressure in laboratory contexts (Dudley and Chai, 1996), we can examine the physiological and aerodynamic consequences of high-altitude flight both separately and in concert.”
The paper relies on this without support.
Air density declines systematically with increased elevation.
“The systematic decline in air density with increased elevation must adversely affect the aerodynamics of hovering flight.”
The paper relies on this without support.
High-altitude hummingbirds often perch while feeding, whereas their lowland counterparts almost never do so.
“Highaltitude hummingbirds, for example, often perch while feeding, whereas their lowland counterparts almost never do so.”
The paper relies on this without support.
Some of the larger hummingbird species are most common at high altitudes.
“Some of the larger species are, somewhat paradoxically, most common at high altitudes, exacerbating the demands of hovering flight.”
The paper relies on this without support.
Ecosystem composition, predators, nutritional resources, and vegetational structure all change dramatically across elevational gradients.
“Clearly, ecosystem composition, predators, nutritional resources, and vegetational structure all change dramatically across elevational gradients.”
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Migrating swans profit from favourable changes in wind conditions at low altitude
Klaassen, M., Beekman, J. H., Kontiokorpi, J., Mulder, R. J. W., & Nolet, B. A. (2004) · Journal of Ornithology · doi:10.1007/s10336-004-0025-x · Cited by 35 · see citing works
Increased capillarity in leg muscle of finches living at altitude
Hepple, R. T., Agey, P. J., Hazelwood, L., Szewczak, J. M., MacMillen, R. E., & Mathieu-Costello, O. (1998) · Journal of Applied Physiology · doi:10.1152/jappl.1998.85.5.1871 · Cited by 34 · see citing works
Altitudinal Variation in Bolivian Birds
Traylor, M. A. (1950) · The Condor · doi:10.2307/1364896 · Cited by 31 · see citing works
Predicting Migratory Flight Altitudes by Physiological Migration Models
Liechti, F., Klaassen, M., & Bruderer, B. (2000) · The Auk · doi:10.1093/auk/117.1.205 · Cited by 31 · see citing works
Collision Between a Vulture and an Aircraft at an Altitude of 37,000 Feet
Laybourne, R. C. (1974) · Digital Commons - University of South Florida (University of South Florida) · Cited by 30 · see citing works
Evaporative and excretory water loss during free flight in pigeons
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Life's Devices: The Physical World of Animals and Plants
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Twenty-Three Testable Predictions About Bird Flight
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Oxygen consumption and power of hovering hummingbirds at varying barometric and oxygen pressures
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Water Balance During Real and Simulated Long-Distance Migratory Flight in the Bar-Tailed Godwit
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THE GIANT HUMMINGBIRD <i>PATAGONA GIGAS</i> IN ECUADOR
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Body temperatures in free-flying pigeons
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Effects of hypobaria on parabronchial gas exchange in normoxic and hypoxic ducks
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Adaptations To Hypoxia In Birds: How To Fly High
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Limits to human locomotor performance: phylogenetic origins and comparative perspectives
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High-Altitude Bird Flight: Exercise in a Hostile Environment
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Hovering Performance of Hummingbirds in Hyperoxic Gas Mixtures
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Self-citationFrigatebirds ride high on thermals
Barbraud, C., Chastel, O., & Weimerskirch, H. (2003) · HAL (Le Centre pour la Communication Scientifique Directe) · Cited by 7 · see citing works
Flagged first, then by citations
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Research works printed but not linked: 1
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Cited by
Sorted by citations
Jankowski, J. E., Londoño, G. A., Robinson, S. K., & Chappell, M. A. (2012) · Ecography · doi:10.1111/j.1600-0587.2012.07785.x · Cited by 251 · see citing works
The trans-Himalayan flights of bar-headed geese ( Anser indicus )
Hawkes, L. A., Balachandran, S., Batbayar, N., Butler, P. J., Frappell, P. B., Milsom, W. K., Natsagdorj, T., Newman, S. H., Scott, G. R., Sathiyaselvam, P., et al. (2011) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1017295108 · Cited by 179 · see citing works
“As a consequence, the partial pressure of oxygen (PO2) in the arterial blood may begin to limit maximum performance (4, 5), although negative effects on the rate of oxygen diffusion may be partially ameliorated by an increase in the gas diffusion coefficient (6).”
BackgroundElevated performance: the unique physiology of birds that fly at high altitudes
Scott, G. R. (2011) · Journal of Experimental Biology · doi:10.1242/jeb.052548 · Cited by 180 · see citing works
“Hypobaria has unique consequences for flying animals, because the mechanical power output needed to sustain lift increases in thin air (Altshuler and Dudley, 2006).”
BackgroundEcophysiology of avian migration in the face of current global hazards
Klaassen, M., Hoye, B. J., Nolet, B. A., & Buttemer, W. A. (2012) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2012.0008 · Cited by 146 · see citing works
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Avian distributions under climate change: towards improved projections
Sorte, F. A. L., & Jetz, W. (2010) · Journal of Experimental Biology · doi:10.1242/jeb.038356 · Cited by 115 · see citing works
“This includes optimal fuel deposition during flight (Alerstam, 1991), energy expenditure, variation in physiology and flight behavior (SchmidtWellenburg et al., 2007), and high-altitude flight performance (Seagram et al., 2001; Altshuler and Dudley, 2006).”
BackgroundGraham, C. H., Parra, J. L., Tinoco, B. A., et al. (2012) · Ecology · doi:10.1890/11-0493.1 · Cited by 108 · see citing works
Zhou, T., Shen, X., Irwin, D. M., et al. (2014) · Mitochondrion · doi:10.1016/j.mito.2014.07.012 · Cited by 92 · see citing works
“Hypoxia is an unavoidable environmental stress for high-altitude birds (Altshuler and Dudley, 2006; Scott, 2011).”
BackgroundHigh-altitude champions: birds that live and migrate at altitude
Laguë, S. L. (2017) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00110.2017 · Cited by 67 · see citing works
“Because of the decreased air density present at high altitudes, generating lift during flight becomes increasingly more difficult (2).”
BackgroundAbiotic conditions shape spatial and temporal morphological variation in North American birds
Youngflesh, C., Saracco, J. F., Siegel, R. B., & Tingley, M. W. (2022) · Nature Ecology & Evolution · doi:10.1038/s41559-022-01893-x · Cited by 84 · see citing works
“Parameter σ789 was modeled as half-normal (normal but with 346 support only over positive values), 347 σ7893 ∼ HN(τ1789 , κ1789), [9] 348”
BackgroundAllometry of hummingbird lifting performance
Altshuler, D. L., Dudley, R., Heredia, S. M., & McGuire, J. A. (2010) · Journal of Experimental Biology · doi:10.1242/jeb.037002 · Cited by 79 · see citing works
“…of hummingbird lifting performance Scaling relationships across elevations Effects of elevational gradients on animal flight performance can be profound and derive from reductions in both air density and oxygen availability (Altshuler and Dudley, 2006; Dillon et al., 2006; Dudley and Chai, 1996).”
BackgroundSelf-citationThe influence of weather on the flight altitude of nocturnal migrants in mid‐latitudes
Kemp, M. U., Shamoun-Baranes, J. Z., Dokter, A. M., van Loon, E. E., & Bouten, W. (2013) · Ibis · doi:10.1111/ibi.12064 · Cited by 82 · see citing works
“…system, theory suggests that atmospheric properties © 2013 British Ornithologists’ Union highly correlated with altitude such as oxygen partial pressure (Altshuler & Dudley 2006) and air density (Pennycuick 2008), along with temperature and humidity, may influence a bird’s flight efficiency.”
BackgroundSurpassing Mt. Everest: extreme flight performance of alpine bumble-bees
Dillon, M. E., & Dudley, R. (2014) · Biology Letters · doi:10.1098/rsbl.2013.0922 · Cited by 75 · see citing works
“Commercially reared Bombus impatiens workers are capable of a 50% increase in vertical force production to lift loads [11], but females have smaller wings relative to body size [2] which could limit flight capacity at extreme elevations [1,2].”
BackgroundShanfelter, A. F., Archambeault, S., & White, M. A. (2019) · Genome Biology and Evolution · doi:10.1093/gbe/evz090 · Cited by 64 · see citing works
Oxygen in demand: How oxygen has shaped vertebrate physiology
Dzal, Y. A., Jenkin, S., Laguë, S. L., Reichert, M. N., York, J. M., & Pamenter, M. E. (2015) · Comparative Biochemistry and Physiology Part A Molecular & Integrative Physiology · doi:10.1016/j.cbpa.2014.10.029 · Cited by 70 · see citing works
“due to hypobaria, which reduces the lift and thrust generated by each wingbeat (Altshuler and Dudley, 2006).”
BackgroundGutiérrez‐Pinto, N., McCracken, K. G., Alza, L., et al. (2014) · Biological Journal of the Linnean Society · doi:10.1111/bij.12249 · Cited by 62 · see citing works
Here comes the sun: multimodal displays are associated with sunlight incidence
Sicsú, P. R., Manica, L. T., Maia, R., et al. (2013) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-013-1574-x · Cited by 57 · see citing works
DuBay, S. G., & Witt, C. C. (2014) · Molecular Ecology · doi:10.1111/mec.12836 · Cited by 49 · see citing works
“At the top of this gradient, animals face hypodense air that increases both the energy requirements for powered flight and the rate of dehydration (Altshuler & Dudley 2006).”
The smallest avian genomes are found in hummingbirds
Gregory, T. R., Andrews, C. B., McGuire, J. A., & Witt, C. C. (2009) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2009.1004 · Cited by 51 · see citing works
Boyce, A. J., Shakya, S. B., Sheldon, F. H., Moyle, R. G., & Martin, T. E. (2019) · The Auk · doi:10.1093/auk/ukz054 · Cited by 52 · see citing works
Pan, S., Zhang, T., Rong, Z., Li, H., Gu, Z., Wu, Q., Dong, S., Liu, Q., Lin, Z., Deutschová, L., et al. (2017) · Molecular Ecology · doi:10.1111/mec.14090 · Cited by 49 · see citing works
“…contribution of hypoxia to MHC class II B gene expression in QH sakers could be little because there is no significant correlation between the MHC class II B gene expression differences and elevation, which is in direct proportion to reduction of oxygen partial pressure (Altshuler & Dudley 2006).”
BackgroundLü, X., Ke, D., Zeng, X., et al. (2009) · Journal of Arid Environments · doi:10.1016/j.jaridenv.2009.06.011 · Cited by 49 · see citing works
Parra, J. L., McGuire, J. A., & Graham, C. H. (2010) · The American Naturalist · doi:10.1086/656619 · Cited by 47 · see citing works
Cadena, C. D., & Loiselle, B. A. (2007) · Ecography · doi:10.1111/j.2007.0906-7590.05045.x · Cited by 47 · see citing works
Mutation, Randomness, and Evolution
Stoltzfus, A. (2021) · doi:10.1093/oso/9780198844457.001.0001 · Cited by 41 · see citing works
Maina, J. N., McCracken, K. G., Chua, B., York, J. M., & Milsom, W. K. (2017) · PLoS ONE · doi:10.1371/journal.pone.0174395 · Cited by 45 · see citing works
Lim, M., Witt, C. C., Graham, C. H., & Dávalos, L. M. (2019) · Genome Biology and Evolution · doi:10.1093/gbe/evz101 · Cited by 42 · see citing works
Flying high: Limits to flight performance by sparrows on the Qinghai-Tibet Plateau
Sun, Y., Ren, Z.-P., Wu, Y., et al. (2016) · Journal of Experimental Biology · doi:10.1242/jeb.142216 · Cited by 43 · see citing works
“Moreover, animal flight at high altitude involves aerodynamic responses to hypodense conditions, along with enhanced oxygen delivery to flight muscles to compensate for its reduced atmospheric partial pressure (Altshuler and Dudley, 2006; Dillon et al., 2006).”
BackgroundThe role of wingbeat frequency and amplitude in flight power
Krishnan, K., Garde, B., Bennison, A., Cole, N. C., Cole, E.-L., Darby, J., Elliott, K. H., Fell, A., Gómez‐Laich, A., de Grissac, S., et al. (2022) · Journal of The Royal Society Interface · doi:10.1098/rsif.2022.0168 · Cited by 34 · see citing works
The role of climate factors in geographic variation in body mass and wing length in a passerine bird
Sun, Y., Li, M., Song, G., et al. (2017) · Avian Research · doi:10.1186/s40657-016-0059-9 · Cited by 37 · see citing works
“The lower wing loadings of longer wings can offset the increased energy requirements incurred by the lower air density at higher altitudes (Swaddle and Lockwood 2003; Altshuler and Dudley 2006).”
Alhajeri, B. H., Schenk, J. J., & Steppan, S. J. (2015) · Biological Journal of the Linnean Society · doi:10.1111/bij.12695 · Cited by 36 · see citing works
“Finally, elevational preference is a major ecological axis on which sister species diverge in various taxa (Endler, 1982; Porter et al., 2002; Navas, 2002; Hall, 2005; Altshuler & Dudley, 2006; Cadena, 2007) and is an important environmental determinant of habitat.”
BackgroundMadsen, P. B., Morabowen, A., Andino, P., Espinosa, R., Cauvy-Fraunié, S., Dangles, O., & Jacobsen, D. (2015) · Ecological Entomology · doi:10.1111/een.12232 · Cited by 34 · see citing works
“…the atmospheric pressure of oxygen decreases with increasing altitude (Dillon et al., 2006), and a low partial pressure of oxygen is recognised as an additional possible limiting factor for life at high altitudes (Halloy, 1989; Navas, 2003; Altshuler & Dudley, 2006; Jankowski et al., 2013).”
BackgroundLee, S. Y., Scott, G. R., & Milsom, W. K. (2008) · Comparative Biochemistry and Physiology Part C Toxicology & Pharmacology · doi:10.1016/j.cbpc.2008.05.009 · Cited by 35 · see citing works
“Indeed, increases in wingbeat amplitude (but not frequency) have been noted in hummingbirds at altitude compared to those at sea level (Altshuler and Dudley, 2006), which could substantially heighten the metabolic costs of flight.”
BackgroundInfluentialKandyel, R., Basyouny, H. A. E., Nahass, E. E., Madkour, F. A., Haddad, S., Massoud, D., Morsy, K., Madkour, N. F., & Abumandour, M. M. A. (2021) · Microscopy Research and Technique · doi:10.1002/jemt.23980 · Cited by 34 · see citing works
Cadena, C. D., & Loiselle, B. A. (2007) · Ecography · doi:10.1111/j.0906-7590.2007.05045.x · Cited by 34 · see citing works
“This assumption has been well-established by functional studies (Porter et al. 2002, Navas 2003, Altshuler and Dudley 2006 ), implying that our approach is closer to characterizing ecological niches than that of studies that only consider elevation, which affects organisms only indirectly.”
ResultInto rude air: hummingbird flight performance in variable aerial environments
Ortega-Jiménez, V. M., Badger, M. A., Wang, H., & Dudley, R. (2016) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2015.0387 · Cited by 32 · see citing works
Herder, E. A., Spence, A. R., Tingley, M. W., & Hird, S. M. (2021) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2020.597756 · Cited by 27 · see citing works
“Hummingbirds are well equipped to deal with most challenges of high elevation, like low ambient temperature, decreased air density, and low oxygen availability (Altshuler and Dudley, 2006).”
BackgroundComparative Energetics of the Giant Hummingbird (Patagona gigas)
Fernández, M. J., Dudley, R., & Bozinovic, F. (2011) · Physiological and Biochemical Zoology · doi:10.1086/660084 · Cited by 29 · see citing works
“Because hummingbird flight performance is progressively challenged at higher elevations (Altshuler and Dudley 2002, 2006), a study of hovering metabolism in P. gigas at different elevations would be of particular interest.”
BackgroundMutation-biased adaptation in Andean house wrens
Stoltzfus, A. B., & McCandlish, D. M. (2015) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1518490112 · Cited by 27 · see citing works
Aeromechanics in aeroecology: flight biology in the aerosphere
Swartz, S. M., Breuer, K., & Willis, D. J. (2007) · Integrative and Comparative Biology · doi:10.1093/icb/icn054 · Cited by 27 · see citing works
“Studies of migratory flight, primarily in birds, but also in insects, do provide some glimpses of the dependence of flight energetics on the conditions of ambient flow (Butler et al. 2000; Weimerskirch et al. 2000; Gannes 2001; Bowlin et al. 2005; Altshuler and Dudley 2006).”
BackgroundThe travel speeds of large animals are limited by their heat-dissipation capacities
Dyer, A., Brose, U., Berti, E., Rosenbaum, B., & Hirt, M. R. (2023) · PLoS Biology · doi:10.1371/journal.pbio.3001820 · Cited by 22 · see citing works
“These behaviours are associated with a lower oxygen availability and an increased mechanical cost of flight at altitude [58,59] or the failure to settle at suitable stopover sites (night flights; [60]) and, therefore, only remain beneficial when heat dissipation is critical to sustained locomotion.”
BackgroundReduced habitat suitability and landscape connectivity in a songbird migratory divide
Justen, H. C., Lee‐Yaw, J. A., & Delmore, K. E. (2021) · Global Ecology and Biogeography · doi:10.1111/geb.13367 · Cited by 22 · see citing works
Pelayo, R. C., Soriano, P. J., Márquez, N., & Navarro, L. (2019) · Plant Ecology & Diversity · doi:10.1080/17550874.2019.1675096 · Cited by 20 · see citing works
“radiation loads and low partial pressures of O2 and CO2 (Altshuler and Dudley 2006; Azócar and Rada 2006; Dillon et al. 2006).”
BackgroundHigh altitude flights by ruddy shelduck Tadorna ferruginea during trans‐Himalayan migrations
Parr, N., Bearhop, S., Douglas, D. C., Takekawa, J. Y., Prosser, D. J., Newman, S. H., Perry, W. M., Balachandran, S., Witt, M. J., Hou, Y. M., et al. (2017) · Journal of Avian Biology · doi:10.1111/jav.01443 · Cited by 21 · see citing works
Byamungu, R. M., Schleuning, M., Ferger, S. W., et al. (2021) · Diversity and Distributions · doi:10.1111/ddi.13403 · Cited by 20 · see citing works
Point: high altitude is for the birds!
Scott, G., Meir, J., Hawkes, L., Frappell, P., & Milsom, W. (2011) · Journal of applied physiology · doi:10.1152/japplphysiol.00821.2011 · Cited by 21 · see citing works
Mechanical Constraints on Flight at High Elevation Decrease Maneuvering Performance of Hummingbirds
Segre, P. S., Dakin, R., Read, T. J. G., Straw, A. D., & Altshuler, D. L. (2016) · Current Biology · doi:10.1016/j.cub.2016.10.028 · Cited by 19 · see citing works
Self-citationParra, J. L., Rahbek, C., McGuire, J. A., & Graham, C. H. (2011) · Journal of Biogeography · doi:10.1111/j.1365-2699.2011.02558.x · Cited by 18 · see citing works
“Because of the energetic demands of flight in hummingbirds at high elevations and the conservatism in flight morphology (Altshuler et al., 2004; Altshuler & Dudley, 2006), we expect all clades to respond to this important axis of environmental variation.”
BackgroundPervasive mimicry in flight behavior among aposematic butterflies
Page, E., Queste, L. M., Rosser, N. S., et al. (2024) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2300886121 · Cited by 18 · see citing works
Pu, Z.-X., & Guo, Y. (2023) · Ecology and Evolution · doi:10.1002/ece3.10492 · Cited by 18 · see citing works
Assessing the accuracy of altitude estimates in avian biologging devices
Lato, K. A., Stepanuk, J. E. F., Heywood, E. I., et al. (2022) · PLoS ONE · doi:10.1371/journal.pone.0276098 · Cited by 18 · see citing works
“For example, many terrestrial birds and seabirds use air thermals for efficient flight and alter their behavior in accordance with air density [53, 54].”
BackgroundBlack Kites on a flyway between Western Siberia and the Indian Subcontinent
Literák, I., Škrábal, J., Karyakin, I. V., Andreyenkova, N. G., & Важов, С. В. (2022) · Scientific Reports · doi:10.1038/s41598-022-09246-1 · Cited by 22 · see citing works
“They y and, moreover, stay for hours resting at night in the environment of mountains at altitudes over 5000 m with variable wind speed and direction, where the air density and partial pressure of oxygen is roughly half of that at sea level (59, 69).”
BackgroundKiat, Y., Davaasuren, B., Erdenechimeg, T., Troupin, D., & Sapir, N. (2020) · Ecography · doi:10.1111/ecog.05287 · Cited by 16 · see citing works
Fog and dense gas disrupt mosquito flight due to increased aerodynamic drag on halteres
Dickerson, A. K., Shankles, P. G., Berry, B. E., et al. (2015) · Journal of Fluids and Structures · doi:10.1016/j.jfluidstructs.2015.03.016 · Cited by 16 · see citing works
“Birds at high altitudes fly with reduced forward drag, but increased requirements for lift production (Altshuler and Dudley, 2006).”
BackgroundParr, N., Bishop, C. M., Batbayar, N., et al. (2019) · Journal of Experimental Biology · doi:10.1242/jeb.203695 · Cited by 15 · see citing works
“Birds that migrate at high altitude may experience all of these: extreme cold, low humidity and reduced oxygen availability as a result of declining air density (Altshuler and Dudley, 2006; Butler, 2010).”
BackgroundMitochondrial DNA study in domestic chicken
Lan, D., Hu, Y., Zhu, Q., & Liu, Y. (2015) · Mitochondrial DNA Part A · doi:10.3109/19401736.2015.1106526 · Cited by 16 · see citing works
“Mitochondrial DNA and Tibetan chicken high-altitude adaptation Hypoxia is an unavoidable environmental stress for high-altitude birds (Altshuler & Dudley, 2006; Scott, 2011).”
BackgroundAdaptations to life at high elevation: An introduction to the symposium
Altshuler, D. L. (2006) · Integrative and Comparative Biology · doi:10.1093/icb/icj002 · Cited by 15 · see citing works
“We took a similar approach for birds that takes into account the additional effects of variable wind speed and humidity, and also examine the consequences of elevational colonization by hummingbirds (Altshuler and Dudley, 2005). The influence of topography in general, and elevation in particular, on avian speciation i…”
BackgroundSelf-citationSander, M. M., & Chamberlain, D. (2020) · Ibis · doi:10.1111/ibi.12843 · Cited by 14 · see citing works
Aerodynamic and dynamic characteristics of flapping wings under low air density
Ha, G., & Park, H. C. (2024) · Acta Astronautica · doi:10.1016/j.actaastro.2024.06.048 · Cited by 12 · see citing works
Liu, Y., Jin, H., Naeem, M., & An, J. (2020) · Apidologie · doi:10.1007/s13592-020-00795-w · Cited by 14 · see citing works
“lower air density and oxygen content than lowaltitude environments and sustain low temperatures (Altshuler and Dudley 2006; Dillon et al. 2006).”
BackgroundColwell, R. K., Rangel, T., Fučíková, K., Sustaita, D., Yanega, G., & Rico-Guevara, A. (2023) · American Naturalist · doi:10.1086/726036 · Cited by 10 · see citing works
Design and Verification of Large-Scaled Flapping Wings for High Altitude Environment
Zhu, Z., Song, B., & Xue, D. (2022) · Applied Sciences · doi:10.3390/app12105140 · Cited by 12 · see citing works
Rivadeneira, G., Ramsay, P. M., & Montúfar, R. (2020) · Alpine Botany · doi:10.1007/s00035-020-00234-7 · Cited by 12 · see citing works
Norman, J. A., Christidis, L., & Schodde, R. (2018) · Organisms Diversity & Evolution · doi:10.1007/s13127-018-0364-8 · Cited by 12 · see citing works
Reardon, S., & Schoeman, M. C. (2017) · Acta Chiropterologica · doi:10.3161/15081109acc2017.19.2.005 · Cited by 12 · see citing works
Upslope migration is slower in insects that depend on metabolically demanding flight
Moore, M. P., Shaich, J., & Stroud, J. T. (2023) · Nature Climate Change · doi:10.1038/s41558-023-01794-2 · Cited by 11 · see citing works
Abiotic conditions shape spatial and temporal morphological variation in North American birdsPreprint
Youngflesh, C., Saracco, J. F., Siegel, R. B., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.02.17.480905 · Cited by 11 · see citing works
Elevational constraints on flight efficiency shape global gradients in avian wing morphology
Yang, J., Yang, C., Lin, H.-W., Lees, A. C., & Tobias, J. A. (2025) · Current Biology · doi:10.1016/j.cub.2025.02.068 · Cited by 9 · see citing works
Mogilicherla, K., CHATTERJEE, R. N., Reddy, M. R., Dange, M., & Bhattacharya, T. K. (2023) · Frontiers in Genetics · doi:10.3389/fgene.2023.1083976 · Cited by 10 · see citing works
“In high-altitude birds, hypoxia is an unavoidable environmental stress; in natural selection, every step of aerobic respiration had to be experienced to improve the adaptability to hypoxia (Altshuler and Dudley, 2006; Scott, 2011; Scott and Milsom, 2006).”
BackgroundDufour, P., Nussbaumer, R., Briedis, M., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.08.18.608464 · Cited by 9 · see citing works
Biliński, T., Bylak, A., Kukuła, K., & Zadrąg‐Tęcza, R. (2021) · PeerJ · doi:10.7717/peerj.12286 · Cited by 9 · see citing works
Clutch and egg size of Tree Swallows along an elevational gradient
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Determining Candidate Hypobaric Hypoxia Profiles for Humane Killing of Laboratory Mice
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“Given their ability for flight at high altitudes, birds have numerous adaptations for enhanced oxygen delivery such as; high hemoglobin oxygen affinity, large lung to blood volume ratios and tubular arrangement of gas exchange systems unlike the spherical alveoli of mammals (36).”
BackgroundFlying High: The Unique Physiology of Birds that Fly at High Altitudes
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Dynamic Temperature Compensation of Pressure Sensors in Migratory Bird Biologging Applications
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Avian Adaptations to High Mountain Habitats
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Sex-dependent elevational effects on bird feather moult
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“We note that several other environmental variables are known to change with elevation, including oxygen partial pressure, which could also possibly impact the physiological tolerance of the animals (Altshuler 2006, Altshuler and Dudley 2006, DuBois et al.”
BackgroundHaas, M., & Janiga, M. (2020) · The European Zoological Journal · doi:10.1080/24750263.2020.1813821 · Cited by 6 · see citing works
On fuel choice and water balance during migratory bird flights.
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The Lesser-Known Transitions: Organismal Form and Function across Abiotic Gradients
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“Longer wings and tails help to increase flight efficiency, maneuverability, as well as balance and lift in conditions of high elevation (Landmann & Winding 1993, Altshuler & Dudley 2006, Maybury & Rayner 2001).”
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Counterpoint: high altitude is not for the birds!
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“Flight speed is negatively related to the square root of the density of air (Hedenstrom and Alerstam 1995; Altshuler and Dudley 2006).”
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“In the Tibetan Plateau, the rarefied air is a typical environmental feature that tests the growth and survival of aerobic organisms (Altschuler and Dudley 2006; Beall 2006; Dillon et al. 2006; West 2006) .”
BackgroundComments on Point:Counterpoint: High altitude is/is not for the birds!
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“Some species of birds even fly nonstop over very long distances while others fly under the hypoxic conditions of extreme altitude [94-98].”
Background
- Authors
- Altshuler, D. L.
- Affiliations
- California Institute of Technology (US)
- Corresponding author
- Altshuler, D. L.
- ORCID
- Altshuler, D. L.
- Venue
- Integrative and Comparative Biology
- Publisher
- Oxford University Press (OUP)
- Published
- 2006-01-06
- Volume
- 46
- Issue
- 1
- Pages
- 62-71
- ISSN
- 1540-7063 (ISSN-L)
- 1557-7023
- OpenAlex
- W2042695519
- Cited by
- 141
- References
- 95
- Citations by year
- 2026: 6 · 2025: 8 · 2024: 7 · 2023: 9 · 2022: 13 · 2021: 31 · 2020: 8 · 2019: 6 · 2018: 5 · 2017: 9 · 2016: 3 · 2015: 8 · 2014: 4 · 2013: 2 · 2012: 3
- FWCI
- 1.52
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Integrative and Comparative Biology (publishedVersion)
- PubMed (publishedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- Funders
- National Science Foundation (0330750, 9817138)
- Earthwatch Institute
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › High Altitude and Hypoxia
- Physical Sciences › Environmental Science › Ecology › Physiological and biochemical adaptations
- Physical Sciences › Engineering › Aerospace Engineering › Biomimetic flight and propulsion mechanisms
- Keywords
- bird flight, high altitude, hypoxia, bird migration, wind speed, air density, oxygen partial pressure, hummingbirds, hovering flight, induced power, wing size, stroke amplitude, flight biomechanics, flight physiology, high-elevation birds
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