WING MORPHOLOGY AND FLIGHT BEHAVIOR OF SOME NORTH AMERICAN HUMMINGBIRD SPECIES

Abstract We explored the relationship between wing morphology and flight behavior with respect to sex and age in five species of North American hummingbirds. We first measured the length, chord or “width,“ and area of entire hummingbird wing planforms. We then calculated additional parameters of wing shape and size, including aspect and shape ratios, degree of taper or “pointedness,“ wing loading, and wing disc loading (WDL). Wings of adult males are not only shorter but also more narrow and tapered than those of adult or immature females; immature males have larger wings and lower WDL, more like those of females. A proposed relationship between WDL and territorial behavior and dominance is not supported, given that adult and immature males show similar feeding territoriality outside the breeding season but females rarely do. The more extreme and divergent wings of adult males probably reflect sexual selection in connection with aerial displays that include species-specific sound effects given during the breeding season. North American species are unusual among hummingbirds in showing reversed sexual size-dimorphism (males smaller, with relatively shorter wings), a feature shared with some other small hummingbirds, notably the “Pygmornis“ hermits. Attempts to explain hummingbird foraging and territorial behavior on the basis of differences in WDL have failed because many aspects of wing morphology, physiology, and flight behavior were not taken into account. Several wing parameters appear more related to other modes of flight than to strategies of nectar exploitation, and the morphology of any given wing represents a compromise between the often conflicting aerodynamic demands of different flight modes. Understanding hummingbird flight will require broad comparative studies of wing morphology and wingbeat kinematics in relation to flight behavior, and new theoretical models and experimental data will be needed to elucidate physiological and aerodynamic mechanisms underlying forward flight and maneuvering. Morfología Alar y Comportamiento de Vuelo de Unas Especies de Colibríes de Norteamérica

[1]  L. Baptista,et al.  The Source of the Dive-Noise of the Anna's Hummingbird , 1979 .

[2]  R. Epting.  Functional Dependence of the Power for Hovering on Wing Disc Loading in Hummingbirds , 1980, Physiological Zoology.

[3]  James H. Brown,et al.  Community Organization in Hummingbirds: Relationships Between Morphology and Ecology , 1985 .

[4]  Black-chinned Hummingbird (Archilochus alexandri) , 2000 .

[5]  Robert Dudley,et al.  Maximum Flight Performance of Hummingbirds: Capacities, Constraints, and Trade‐Offs , 1999, The American Naturalist.

[6]  P. Feinsinger,et al.  On the Relationship between Wing Disc Loading and Foraging Strategy in Hummingbirds , 1975, The American Naturalist.

[7]  A. Zahavi.  Mate selection-a selection for a handicap. , 1975, Journal of theoretical biology.

[8]  R. W. Russell,et al.  Exploitative compensation by subordinate age-sex classes of migrant rufous hummingbirds , 1993, Behavioral Ecology and Sociobiology.

[9]  R. M. Alexander,et al.  Elastic mechanisms in animal movement , 1988 .

[10]  F. G. Stiles.  Behavioral, Ecological and Morphological Correlates of Foraging for Arthropods by the Hummingbirds of a Tropical Wet Forest , 1995 .

[11]  Pilar A. Vásquez,et al.  THE COURTSHIP DISPLAYS AND SOCIAL SYSTEM OF THE WHITE-RUFFED MANAKIN IN COSTA RICA , 2002 .

[12]  Robert Dudley,et al.  The ecological and evolutionary interface of hummingbird flight physiology. , 2002, The Journal of experimental biology.

[13]  Douglas L. Altshuler,et al.  Aerodynamic forces of revolving hummingbird wings and wing models , 2004 .

[14]  Scott Weidensaul,et al.  Ruby-throated Hummingbird (Archilochus colubris) , 2019, Birds of the World.

[15]  U. Norberg.  Vertebrate Flight: Mechanics, Physiology, Morphology, Ecology and Evolution , 1990 .

[16]  T. Casey.  A Comparison of Mechanical and Energetic Estimates of Flight Cost for Hovering Sphinx Moths , 1981 .

[17]  D. Inouye,et al.  Roles of the wing whistle in the territorial behaviour of male broad-tailed hummingbirds (Selasphorus platycercus) , 1983, Animal Behaviour.

[18]  R. W. Russell,et al.  Interference asymmetries among age-sex classes of rufous hummingbirds during migratory stopovers , 1993, Behavioral Ecology and Sociobiology.

[19]  F. G. Stiles,et al.  GEOGRAPHICAL ASPECTS OF BIRD-FLOWER COEVOLUTION, WITH PARTICULAR REFERENCE TO CENTRAL AMERICA' , 1981 .

[20]  Robert K. Colwell,et al.  Community Organization Among Neotropical Nectar-Feeding Birds , 1978 .

[21]  D. Altshuler,et al.  Hovering performance of hummingbirds in hyperoxic gas mixtures. , 2001, The Journal of experimental biology.

[22]  F. A. Pitelka.  Territoriality and Related Problems in North American Hummingbirds , 1942 .

[23]  R. Dudley,et al.  Of Hummingbirds and Helicopters: Hovering Costs, Competitive Ability, and Foraging Strategies , 2004, The American Naturalist.

[24]  R Dudley,et al.  Transient hovering performance of hummingbirds under conditions of maximal loading. , 1997, The Journal of experimental biology.

[25]  R. K. Colwell.  Rensch’s Rule Crosses the Line: Convergent Allometry of Sexual Size Dimorphism in Hummingbirds and Flower Mites , 2000, The American Naturalist.

[26]  T. Weis-Fogh.  Energetics of Hovering Flight in Hummingbirds and in Drosophila , 1972 .

[27]  D. Altshuler,et al.  Ecophysiology of hummingbird flight along elevational gradients: an integrated approach , 2001 .

[28]  D. Altshuler,et al.  CONFLICTING TERMINOLOGY FOR WING MEASUREMENTS IN ORNITHOLOGY AND AERODYNAMICS , 2004 .

[29]  F. G. Stiles.  Food supply and the annual cycle of the Anna hummingbird , 1973 .

[30]  James H. Brown,et al.  Influence of Economics, Interspecific Competition, and Sexual Dimorphism on Territoriality of Migrant Rufous Hummingbirds , 1978 .

[31]  M. Hixon,et al.  Weight gain and adjustment of feeding territory size in migrant hummingbirds. , 1983, Proceedings of the National Academy of Sciences of the United States of America.

[32]  P. Ewald.  Influence of asymmetries in resource quality and age on aggression and dominance in black-chinned hummingbirds , 1985, Animal Behaviour.

[33]  R. Dudley.  The Biomechanics of Insect Flight: Form, Function, Evolution , 1999 .

[34]  W. Calder,et al.  The hummingbird's restraint: A natural model for weight control , 1990, Experientia.

[35]  F. G. Stiles.  A New Species of Emerald Hummingbird (Trochilidae, Chlorostilbon) from the Sierra de Chiribiquete, Southeastern Colombia, with a Review of the C. mellisugus Complex , 1996 .

[36]  C. Gans,et al.  Variations in the distribution of motor end‐plates in the avian pectoralis , 1993, Journal of morphology.

[37]  R. S. Mulvihill,et al.  A possible relationship between reversed sexual size dimorphism and reduced male survivorship in the ruby-throated hummingbird , 1992 .

[38]  C. Ellington.  The Aerodynamics of Hovering Insect Flight. VI. Lift and Power Requirements , 1984 .

[39]  J. Rayner.  A vortex theory of animal flight. Part 2. The forward flight of birds , 1979, Journal of Fluid Mechanics.

[40]  F. Hainsworth.  Foraging Efficiency and Parental Care in Colibri coruscans , 1977 .

[41]  F. G. Stiles.  Aggressive and Courtship Displays of the Male Anna's Hummingbird , 1982 .

[42]  T. Casey,et al.  Power Output and Wing Disc Loading in Hovering Hummingbirds , 1973, The American Naturalist.

[43]  C. Ellington.  THE AERODYNAMICS OF HOVERING INSECT FLIGHT. V. A VORTEX THEORY , 1984 .

[44]  R. Bleiweiss,et al.  DNA hybridization evidence for the principal lineages of hummingbirds (Aves:Trochilidae). , 1997, Molecular biology and evolution.

[45]  T. Weis-Fogh.  Quick estimates of flight fitness in hovering animals , 1973 .

[46]  J. Usherwood,et al.  The aerodynamics of revolving wings I. Model hawkmoth wings. , 2002, The Journal of experimental biology.