Kenneth P. Dial

Bio: Kenneth P. Dial is an academic researcher from University of Montana. The author has contributed to research in topics: Wing & Wing-assisted incline running. The author has an hindex of 41, co-authored 82 publications receiving 4631 citations. Previous affiliations of Kenneth P. Dial include Northern Arizona University & Harvard University.

Flight kinematics of black-billed magpies and pigeons over a wide range of speeds

Abstract: To investigate how birds that differ in morphology change their wing and body movements while flying at a range of speeds, we analyzed high-speed (60 Hz) video tapes of black-billed magpies (Pica pica) flying at speeds of 4-14 m s-1 and pigeons (Columba livia) flying at 6-20 m s-1 in a wind-tunnel. Pigeons had higher wing loading and higher-aspect-ratio wings compared with magpies. Both species alternated phases of steady-speed flight with phases of acceleration and deceleration, particularly at intermediate flight speeds. The birds modulated their wingbeat kinematics among these phases and frequently exhibited non-flapping phases while decelerating. Such modulation in kinematics during forward flight is typical of magpies but not of pigeons in the wild. The behavior of the pigeons may have been a response to the reduced power costs for flight in the closed wind-tunnel relative to those for free flight at similar speeds. During steady-speed flight, wingbeat frequency did not change appreciably with increasing flight speed. Body angle relative to the horizontal, the stroke-plane angles of the wingtip and wrist relative to the horizontal and the angle describing tail spread at mid-downstroke all decreased with increasing flight speed, thereby illustrating a shift in the dominant function of wing flapping from weight support at slow speeds to positive thrust at fast speeds. Using wingbeat kinematics to infer lift production, it appeared that magpies used a vortex-ring gait during steady-speed flight at all speeds whereas pigeons used a vortex-ring gait at 6 and 8 m s-1, a transitional vortex-ring gait at 10 m s-1, and a continuous-vortex gait at faster speeds. Both species used a vortex-ring gait for acceleration and a continuous-vortex gait or a non-flapping phase for deceleration during flight at intermediate wind-tunnel speeds. Pigeons progressively flexed their wings during glides as flight speed increased but never performed bounds. Wingspan during glides in magpies did not vary with flight speed, but the percentage of bounds among non-flapping intervals increased with speed from 10 to 14 m s-1. The use of non-flapping wing postures seemed to be related to the gaits used during flapping and to the aspect ratio of the wings. We develop an 'adverse-scaling' hypothesis in which it is proposed that the ability to reduce metabolic and mechanical power output using flap-bounding flight at fast flight speeds is scaled negatively with body mass. This represents an alternative to the 'fixed-gear' hypothesis previously suggested by other authors to explain the use of intermittent flight in birds. Future comparative studies in the field would be worthwhile, especially if instantaneous flight speeds and within-wingbeat kinematics were documented; new studies in the laboratory should involve simultaneous recording of wing kinematics and aerodynamic forces on the wing.

Locomotor modules and the evolution of avian flight.

Abstract: The evolution of avian flight can be interpreted by analyzing the sequence of modifications of the primitive tetrapod locomotor system through time. Herein, we introduce the term "locomotor module" to identify anatomical subregions of the musculoskeletal system that are highly integrated and act as functional units during locomotion. The first tetrapods, which employed lateral undulations of the entire body and appendages, had one large locomotor module. Basal dinosaurs and theropods were bipedal and possessed a smaller locomotor module consisting of the hind limb and tail. Bird flight evolved as the superimposition of a second (aerial) locomotor capability onto the ancestral (terrestrial) theropod body plan. Although the origin of the wing module was the primary innovation, alterations in the terrestrial system were also significant. We propose that the primitive theropod locomotor module was functionally and anatomically subdivided into separate pelvic and caudal locomotor modules. This decoupling freed the tail to attain a new and intimate affiliation with the forelimb during flight, a configuration unique to birds. Thus, the evolution of flight can be viewed as the origin and novel association of locomotor modules. Differential elaboration of these modules in various lineages has produced the diverse locomotor abilities of modern birds.

Comparative power curves in bird flight

Abstract: The relationship between mechanical power output and forward velocity in bird flight is controversial, bearing on the comparative physiology and ecology of locomotion1,2. Applied to flying birds, aerodynamic theory predicts that mechanical power should vary as a function of forward velocity in a U-shaped curve. The only empirical test of this theory, using the black-billed magpie (Pica pica), suggests that the mechanical power curve is relatively flat over intermediate velocities3. Here, by integrating in vivo measurements of pectoralis force and length change with quasi-steady aerodynamic models developed using data on wing and body movement, we present mechanical power curves for cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria). In contrast to the curve reported for magpies3, the power curve for cockatiels is acutely concave, whereas that for doves is intermediate in shape and shows higher mass-specific power output at most speeds. We also find that wing-beat frequency and mechanical power output do not necessarily share minima in flying birds. Thus, aspects of morphology, wing kinematics and overall style of flight can greatly affect the magnitude and shape of a species' power curve.

Wing-assisted incline running and the evolution of flight.

Abstract: Flapping wings of galliform birds are routinely used to produce aerodynamic forces oriented toward the substrate to enhance hindlimb traction. Here, I document this behavior in natural and laboratory settings. Adult birds fully capable of aerial flight preferentially employ wing-assisted incline running (WAIR), rather than flying, to reach elevated refuges (such as cliffs, trees, and boulders). From the day of hatching and before attaining sustained aerial flight, developing ground birds use WAIR to enhance their locomotor performance through improved foot traction, ultimately permitting vertical running. WAIR provides insight from behaviors observable in living birds into the possible role of incipient wings in feathered theropod dinosaurs and offers a previously unstudied explanation for the evolution of avian flight.

Allometry of behavior

Abstract: The study of allometric and size scaling relationships is well developed in most biological fields, but lags behind in the area of animal behavior. Part of the reason for this deficit is that scaling relationships of behaviors tend to be inherently more 'noisy' than other biological scaling relationships. However, body size has a pervasive influence on the performance of animals in their environments. For example, the frequently strong relationship between power-to-mass ratios and locomotor performance means that smaller species and individuals enjoy superior locomotor performance (burst acceleration and maneuverability) than larger species, particularly within a clade. We suggest that these size-related functional influences on performance profoundly influence many aspects of animal behavior, such as how animals forage, fight, flee, perceive danger, respond to risk and interact with other individuals. We outline exciting avenues for research on the allometry of behavior by integrating scaling and functional perspectives.

The Theory of Island Biogeography

Abstract: Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

Testing for phylogenetic signal in comparative data: behavioral traits are more labile.

Abstract: The primary rationale for the use of phylogenetically based statistical methods is that phylogenetic signal, the tendency for related species to resemble each other, is ubiquitous. Whether this assertion is true for a given trait in a given lineage is an empirical question, but general tools for detecting and quantifying phylogenetic signal are inadequately developed. We present new methods for continuous-valued characters that can be implemented with either phylogenetically independent contrasts or generalized least-squares models. First, a simple randomization procedure allows one to test the null hypothesis of no pattern of similarity among relatives. The test demonstrates correct Type I error rate at a nominal α = 0.05 and good power (0.8) for simulated datasets with 20 or more species. Second, we derive a descriptive statistic, K, which allows valid comparisons of the amount of phylogenetic signal across traits and trees. Third, we provide two biologically motivated branch-length transformat...

How Many Species Are There on Earth and in the Ocean

Abstract: The diversity of life is one of the most striking aspects of our planet; hence knowing how many species inhabit Earth is among the most fundamental questions in science. Yet the answer to this question remains enigmatic, as efforts to sample the world's biodiversity to date have been limited and thus have precluded direct quantification of global species richness, and because indirect estimates rely on assumptions that have proven highly controversial. Here we show that the higher taxonomic classification of species (i.e., the assignment of species to phylum, class, order, family, and genus) follows a consistent and predictable pattern from which the total number of species in a taxonomic group can be estimated. This approach was validated against well-known taxa, and when applied to all domains of life, it predicts ∼8.7 million (±1.3 million SE) eukaryotic species globally, of which ∼2.2 million (±0.18 million SE) are marine. In spite of 250 years of taxonomic classification and over 1.2 million species already catalogued in a central database, our results suggest that some 86% of existing species on Earth and 91% of species in the ocean still await description. Renewed interest in further exploration and taxonomy is required if this significant gap in our knowledge of life on Earth is to be closed.

How Animals Move: An Integrative View

Abstract: Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.

Getting the measure of biodiversity

Abstract: The term 'biodiversity' is a simple contraction of 'biological diversity', and at first sight the concept is simple too: biodiversity is the sum total of all biotic variation from the level of genes to ecosystems. The challenge comes in measuring such a broad concept in ways that are useful. We show that, although biodiversity can never be fully captured by a single number, study of particular facets has led to rapid, exciting and sometimes alarming discoveries. Phylogenetic and temporal analyses are shedding light on the ecological and evolutionary processes that have shaped current biodiversity. There is no doubt that humans are now destroying this diversity at an alarming rate. A vital question now being tackled is how badly this loss affects ecosystem functioning. Although current research efforts are impressive, they are tiny in comparison to the amount of unknown diversity and the urgency and importance of the task.