Sampling affects population genetic inference: A case study of the Allen's (Selasphorus sasin) and rufous hummingbird (Selasphorus rufus).

Gene flow can affect evolutionary inference when species are undersampled. Here, we evaluate the effects of gene flow and geographic sampling on demographic inference of 2 hummingbirds that hybridize, Allen's hummingbird (Selasphorus sasin) and rufous hummingbird (Selasphorus rufus). Using whole-genome data and extensive geographic sampling, we find widespread connectivity, with introgression far beyond the Allen's × rufous hybrid zone, although the Z chromosome resists introgression beyond the hybrid zone. We test alternative hypotheses of speciation history of Allen's, rufous, and Calliope (S. calliope) hummingbird and find that rufous hummingbird is the sister taxon to Allen's hummingbird, and Calliope hummingbird is the outgroup. A model treating the 2 subspecies of Allen's hummingbird as a single panmictic population fit observed genetic data better than models treating the subspecies as distinct populations, in contrast to morphological and behavioral differences and analyses of spatial population structure. With additional sampling, our study builds upon recent studies that came to conflicting conclusions regarding the evolutionary histories of these 2 species. Our results stress the importance of thorough geographic sampling when assessing demographic history in the presence of gene flow.

The Inverse Krogh Principle: All Organisms Are Worthy of Study.

AbstractKrogh's principle states, "For such a large number of problems there will be some animal of choice, or a few such animals, on which it can be most conveniently studied." The downside of picking a question first and then finding an ideal organism on which to study it is that it will inevitably leave many organisms neglected. Here, we promote the inverse Krogh principle: all organisms are worthy of study. The inverse Krogh principle and the Krogh principle are not opposites. Rather, the inverse Krogh principle emphasizes a different starting point for research: start with a biological unit, such as an organism, clade, or specific organism trait, then seek or create tractable research questions. Even the hardest-to-study species have research questions that can be asked of them: Where does it fall within the tree of life? What resources does it need to survive and reproduce? How does it differ from close relatives? Does it have unique adaptations? The Krogh and inverse Krogh approaches are complementary, and many research programs naturally include both. Other considerations for picking a study species include extreme species, species informative for phylogenetic analyses, and the creation of models when a suitable species does not exist. The inverse Krogh principle also has pitfalls. A scientist that picks the organism first might choose a research question not really suited to the organism, and funding agencies rarely fund organism-centered grant proposals. The inverse Krogh principle does not call for all organisms to receive the same amount of research attention. As knowledge continues to accumulate, some organisms-models-will inevitably have more known about them than others. Rather, it urges a broader search across organismal diversity to find sources of inspiration for research questions and the motivation needed to pursue them.

Great Gray Owls hunting voles under snow hover to defeat an acoustic mirage.

How do Great Gray Owls (Strix nebulosa) capture voles (Cricetidae) through a layer of snow? As snow is a visual barrier, the owls locate voles by ear alone. To test how snow absorbs and refracts vole sound, we inserted a loudspeaker under the snowpack and analysed sound from the loudspeaker, first buried, then unburied. Snow attenuation coefficients rose with frequency (0.3 dB cm-1 at 500 Hz, 0.6 dB cm-1 at 3 kHz) such that low-frequency sound transmitted best. The Great Gray Owl has the largest facial disc of any owl, suggesting they are adapted to use this low-frequency sound. We used an acoustic camera to spatially localize sound source location, and show that snow also refracts prey sounds (refractive index: 1.16). To an owl not directly above the prey, this refraction creates an 'acoustic mirage': prey acoustic position is offset from its actual location. Their hunting strategy defeats this mirage because they hover directly over prey, which is the listening position with least refraction and least attenuation. Among all birds, the Great Gray Owl has the most extreme wing morphologies associated with quiet flight. These extreme wing traits may function to reduce the sounds of hovering, with implications for bioinspiration.

Oh, snap! A within-wing sonation in black-tailed trainbearers.

Vertebrates communicate through a wide variety of sounds, but few mechanisms of sound production, besides vocalization, are well understood. During high-speed dives, male trainbearer hummingbirds (Lesbia spp.) produce a repeated series of loud snaps. Hypotheses for these peculiar sounds include the birds employing their elongated tails and/or striking their wings against each other. Each snap to human ears seems like a single acoustic event, but sound recordings revealed that each snap is actually a couplet of impulsive, atonal sounds produced ∼13 ms apart. Analysis of high-speed videos refutes these previous hypotheses, and furthermore suggests that this sonation is produced by a within-wing mechanism - each instance of a sound coincided with a distinctive pair of deep wingbeats (with greater stroke amplitude, measured for one display sequence). Across many displays, we found a tight alignment between a pair of stereotyped deep wingbeats (in contrast to shallower flaps across the rest of the dive) and patterns of snap production, evidencing a 1:1 match between these sonations and stereotyped kinematics. Other birds including owls and poorwills are reported to produce similar sounds, suggesting that this mechanism of sound production could be somewhat common within birds, yet its physical acoustics remain poorly understood.

Ways that Animal Wings Produce Sound.

There are at least eight ways that wings potentially produce sound. Five mechanisms are aerodynamic sounds, created by airflow, and three are structural sound created by interactions of solid surfaces. Animal flight is low Mach (M), meaning all animals move at <30% of the speed of sound. Thus in aerodynamic mechanisms the effects of air compressibility can be ignored, except in mechanism #1. Mechanism #1 is trapped air, in which air approaches or exceeds Mach 1 as it escapes a constriction. This mechanism is hypothetical but likely. #2 is Gutin sound, the aerodynamic reaction to lift and drag. This mechanism is ubiquitous in flight, and generates low frequency sound such as the humming of hummingbirds or insect wing tones. #3 is turbulence-generated atonal whooshing sounds, which are also widespread in animal flight. #4 are whistles, tonal sounds generated by geometry-induced flow feedback. This mechanism is hypothetical. #5 is aeroelastic flutter, sound generated by elasticity-induced feedback that is usually but not always tonal. This is widespread in birds (feathers are predisposed to flutter) but apparently not bats or insects. Mechanism #6 is rubbing sound (including stridulation), created when bird feathers or insect wings slide past each other. Atonal rubbing sounds are widespread in bird flight and insects; tonal stridulation is widespread in insects. #7 is percussion, created when two stiff elements collide and vibrate, and is present in some birds and insects. Mechanism #8 are tymbals and other bistable conformations. These are stiff elements that snap back and forth between two conformations, producing impulsive, atonal sound. Tymbals are widespread in insects but not birds or bats; insect cuticle appears predisposed to form tymbals. There are few examples of bat wing sounds: are bats intrinsically quiet, or just under-studied? These mechanisms, especially Gutin sound, whooshes, and rubbing (#2, #3, and #6) are prominent cues in ordinary flight of all flying animals, and are the "acoustic substrate" available to be converted from an adventitious sound (cue) into a communication signal. For instance, wing sounds have many times evolved into signals that are incorporated into courtship displays. Conversely, these are the sounds selected to be suppressed if quiet flight is selected for. The physical mechanisms that underlie animal sounds provide context for understanding the ways in which signals and cues may evolve.

The population genetics of nonmigratory Allen's Hummingbird (Selasphorus sasin sedentarius) following a recent mainland colonization.

Allen's Hummingbird comprises two subspecies, one migratory (Selasphorus sasin sasin) and one nonmigratory (S. s. sedentarius). The nonmigratory subspecies, previously endemic to the California Channel Islands, apparently colonized the California mainland on the Palos Verdes Peninsula some time before 1970 and now breeds throughout coastal southern California. We sequenced and compared populations of mainland nonmigratory Allen's Hummingbird to Channel Island populations from Santa Catalina, San Clemente, and Santa Cruz Island. We found no evidence of founder effects on the mainland population. Values of nucleotide diversity on the mainland were higher than on the Channel Islands. There were low levels of divergence between the Channel Islands and the mainland, and Santa Cruz Island was the most genetically distinct. Ecological niche models showed that rainfall and temperature variables on the Channel Islands are similar in the Los Angeles basin and predicted continued expansion of nonmigratory Allen's Hummingbird north along the coast and inland. We also reviewed previous genetic studies of vertebrate species found on the Channel Islands and mainland and showed that broad conclusions regarding island-mainland patterns remain elusive. Challenges include the idiosyncratic nature of colonization itself as well as the lack of a comprehensive approach that incorporates similar markers and sampling strategies across taxa, which, within the context of a comparative study of island-mainland relationships, may lead to inconsistent results.

Introduction to the Symposium: Bio-Inspiration of Quiet Flight of Owls and Other Flying Animals: Recent Advances and Unanswered Questions.

Animal wings produce an acoustic signature in flight. Many owls are able to suppress this noise to fly quietly relative to other birds. Instead of silent flight, certain birds have conversely evolved to produce extra sound with their wings for communication. The papers in this symposium synthesize ongoing research in "animal aeroacoustics": the study of how animal flight produces an acoustic signature, its biological context, and possible bio-inspired engineering applications. Three papers present research on flycatchers and doves, highlighting work that continues to uncover new physical mechanisms by which bird wings can make communication sounds. Quiet flight evolves in the context of a predator-prey interaction, either to help predators such as owls hear its prey better, or to prevent the prey from hearing the approaching predator. Two papers present work on hearing in owls and insect prey. Additional papers focus on the sounds produced by wings during flight, and on the fluid mechanics of force production by flapping wings. For instance, there is evidence that birds such as nightbirds, hawks, or falcons may also have quiet flight. Bat flight appears to be quieter than bird flight, for reasons that are not fully explored. Several research avenues remain open, including the role of flapping versus gliding flight or the physical acoustic mechanisms by which flight sounds are reduced. The convergent interest of the biology and engineering communities on quiet owl flight comes at a time of nascent developments in the energy and transportation sectors, where noise and its perception are formidable obstacles.

Humming hummingbirds, insect flight tones and a model of animal flight sound.

Why do hummingbirds hum and insects whine when their wings flap in flight? Gutin proposed that a spinning propeller produces tonal sound because the location of the center of aerodynamic pressure on each blade oscillates relative to an external receiver. Animal wings also move, and in addition, aerodynamic force produced by animal wings fluctuates in magnitude and direction over the course of the wingbeat. Here, we modeled animal wing tone as the equal, opposite reaction to aerodynamic forces on the wing, using Lowson's equation for the sound field produced by a moving point force. Two assumptions of Lowson's equation were met: animal flight is low (<0.3) Mach and animals from albatrosses to mosquitoes are acoustically compact, meaning they have a small spatial extent relative to the wavelength of their wingbeat frequency. This model predicted the acoustic waveform of a hovering Costa's hummingbird (Calypte costae), which varies in the x, y and z directions around the animal. We modeled the wing forces of a hovering animal as a sinusoid with an amplitude equal to body weight. This model predicted wing sound pressure levels below a hovering hummingbird and mosquito to within 2 dB; and that far-field mosquito wing tone attenuates to 20 dB within about 0.2 m of the animal, while hummingbird humming attenuates to 20 dB at about 10 m. Wing tone plays a role in communication of certain insects, such as mosquitoes, and influences predator-prey interactions, because it potentially reveals the predator's presence to its intended prey.

Sonations in Migratory and Non-migratory Fork-tailed Flycatchers (Tyrannus savana).

Sonations are sounds that animals produce with structures other than the vocal apparatus for communication. In birds, many sonations are usually produced with modified flight feathers through diverse kinematic mechanisms. For instance, aeroelastic fluttering of feathers produces tonal sound when airflow exceeds a threshold velocity and induces flight feathers to oscillate at a constant frequency. The Fork-tailed flycatcher (Tyrannus savana) is a Neotropical bird with both migratory and year-round resident subspecies that differ in the shape of the outer primary feathers of their wings. By integrating behavioral observations, audio recordings, and high-speed videos, we find that male Fork-tailed flycatchers produce sonations with their outer primary feathers P8-10, and possibly P7. These sounds are produced during different behavioral contexts including: the pre-dawn display, intraspecific territorial disputes, when attacking potential nest predators, and when escaping. By placing feathers in a wind tunnel, we elicited flutter at frequencies that matched the acoustic signature of sounds recorded in the wild, indicating that the kinematic mechanism responsible for sound production is aeroelastic flutter. Video of wild birds indicated that sonations were produced during the downstroke. Finally, the feathers of migratory (T.s.savana) and year-round resident (T.s.monachus) Fork-tailed flycatchers flutter in feather locations that differ in shape between the subspecies, and these shape differences between the subspecies result in sounds produced at different frequencies.

Evidence that the Dorsal Velvet of Barn Owl Wing Feathers Decreases Rubbing Sounds during Flapping Flight.

Owls have specialized feather features hypothesized to reduce sound produced during flight. One of these features is the velvet, a structure composed of elongated filaments termed pennulae that project dorsally from the upper surface of wing and tail feathers. There are two hypotheses of how the velvet functions to reduce sound. According to the aerodynamic noise hypothesis, the velvet reduces sound produced by aerodynamic processes, such as turbulence development on the surface of the wing. Alternatively, under the structural noise hypothesis, the velvet reduces frictional noise produced when two feathers rub together. The aerodynamic noise hypothesis predicts impairing the velvet will increase aerodynamic flight sounds predominantly at low frequency, since turbulence formation predominantly generates low frequency sound; and that changes in sound levels will occur predominantly during the downstroke, when aerodynamic forces are greatest. Conversely, the frictional noise hypothesis predicts impairing the velvet will cause a broadband (i.e., across all frequencies) increase in flight sounds, since frictional sounds are broadband; and that changes in sound levels will occur during the upstroke, when the wing feathers rub against each other the most. Here, we tested these hypotheses by impairing with hairspray the velvet on inner wing feathers (P1-S4) of 13 live barn owls (Tyto alba) and measuring the sound produced between 0.1 and 16 kHz during flapping flight. Relative to control flights, impairing the velvet increased sound produced across the entire frequency range (i.e., the effect was broadband) and the upstroke increased more than the downstroke, such that the upstroke of manipulated birds was louder than the downstroke, supporting the frictional noise hypothesis. Our results suggest that a substantial amount of bird flight sound is produced by feathers rubbing against feathers during flapping flight.

Specialized Feathers Produce Sonations During Flight in Columbina Ground Doves.

The shape of remiges (primary and secondary feathers) is constrained and stereotyped by the demands of flight, but members of the subfamily of New World ground doves (Peristerinae) possess many atypical remex shapes with which they produce sonations of alarm. Within the genus Columbina specifically, the seventh primary feathers (P7) have elongated barbs that create a protrusion on the trailing vane which varies in size and shape between species. These feathers are hypothesized to have been coopted to produce communicative sounds (i.e., sonations) during flight, but the mechanism of this sound production is unknown. We tested the sound-producing capabilities of spread wing specimens from three species of ground doves (C. inca, C. passerina, and C. talpacoti) in a wind tunnel. High speed video and audio analyses indicated that all wings of adult birds produced buzzing sounds in the orientation and flow velocity of mid-upstroke. These buzzing sounds were produced as the protrusion of elongated barbs fluttered and collided with adjacent P6 feathers at a fundamental frequency of 200 and 400 Hz, respectively. Wings from juvenile C. inca produced significantly quieter buzzes and most (three of four individuals) lacked the elongated barbs that are present in adults. Buzzing sounds produced in the wind tunnel were similar to those produced by wild birds indicating that these P7 feathers have been coopted to produce acoustic signals (sonations) during flight. The shape and mechanism of sound production described here in Columbina appear to be unique among birds.

Evolutionary and Ecological Correlates of Quiet Flight in Nightbirds, Hawks, Falcons, and Owls.

Two hypotheses have been proposed for the evolution of structures that reduce flight sounds in birds. According to the stealth hypothesis, flying quietly reduces the ability of other animals (e.g., prey) to detect the animal's approach from its flight sounds. This hypothesis predicts that animals hunting prey with acute hearing evolve silencing features. The self-masking hypothesis posits that reduced flight sounds permit the animal itself to hear better (such as the sounds of its prey, or its own echolocation calls) during flight. This hypothesis predicts that quieting features evolve in predators that hunt by ear, or in species that echolocate. Owls, certain hawks, and nightbirds (nocturnal Caprimulgiformes) have convergently evolved a sound-reducing feature: a velvety coating on the dorsal surface of wing and tail feathers. Here we document a fourth independent origin of the velvet, in the American kestrel (Falco sparverius). Among these four clades (hawks, falcons, nightbirds, and owls), the velvet is longer and better developed in wing and tail regions prone to rubbing with neighboring feathers, apparently to reduce broadband frictional noise produced by rubbing of adjacent feathers. We tested whether stealth or self-masking better predicted which species evolved the velvet. There was no support of echolocation as a driver of the velvet: oilbird(Steatornis caripensis) and glossy swiftlet (Collocalia esculenta) each evolved echolocation but neither had any velvet. A phylogenetic least squares fit of stealth and self-masking (to better hear prey sounds) provided support for both hypotheses. Some nightbirds (nightjars, potoos, and owlet-nightjars) eat flying insects that do not make much sound, implying the velvet permits stealthy approach of flying insects. One nightbird clade, frogmouths (Podargus) have more extensive velvet than other nightbirds and may hunt terrestrial prey by ear, in support of self-masking. In hawks, the velvet is also best developed in species known or suspected to hunt by ear (harriers and kites), supporting the self-masking hypothesis, but velvet is also present in reduced form in hawk species not known to hunt by ear, in support of the stealth hypothesis. American kestrel is not known to hunt by ear, and unlike the other falcons sampled, flies slowly (kite-like) when hunting. Thus the presence of velvet in it supports the stealth hypothesis. All owls sampled (n = 13 species) had extensive velvet, including the buffy fish-owl (Ketupa ketupu), contrary to literature claims that fish-owls had lost the velvet. Collectively, there is support for both the self-masking and stealth hypotheses for the evolution of dorsal velvet in birds.

Evolution and Ecology of Silent Flight in Owls and Other Flying Vertebrates.

We raise and explore possible answers to three questions about the evolution and ecology of silent flight of owls: (1) do owls fly silently for stealth, or is it to reduce self-masking? Current evidence slightly favors the self-masking hypothesis, but this question remains unsettled. (2) Two of the derived wing features that apparently evolved to suppress flight sound are the vane fringes and dorsal velvet of owl wing feathers. Do these two features suppress aerodynamic noise (sounds generated by airflow), or do they instead reduce structural noise, such as frictional sounds of feathers rubbing during flight? The aerodynamic noise hypothesis lacks empirical support. Several lines of evidence instead support the hypothesis that the velvet and fringe reduce frictional sound, including: the anatomical location of the fringe and velvet, which is best developed in wing and tail regions prone to rubbing, rather than in areas exposed to airflow; the acoustic signature of rubbing, which is broadband and includes ultrasound, is present in the flight of other birds but not owls; and the apparent relationship between the velvet and friction barbules found on the remiges of other birds. (3) Have other animals also evolved silent flight? Wing features in nightbirds (nocturnal members of Caprimulgiformes) suggest that they may have independently evolved to fly in relative silence, as have more than one diurnal hawk (Accipitriformes). We hypothesize that bird flight is noisy because wing feathers are intrinsically predisposed to rub and make frictional noise. This hypothesis suggests a new perspective: rather than regarding owls as silent, perhaps it is bird flight that is loud. This implies that bats may be an overlooked model for silent flight. Owl flight may not be the best (and certainly, not the only) model for "bio-inspiration" of silent flight.

Proponemos y exploramos posibles respuestas a tres preguntas sobre la evolución y ecología del vuelo silencioso en lechuzas: (1) ¿Las lechuzas vuelan silenciosamente por sigilo o para reducir el auto-enmascaramiento?. La evidencia actual favorece levemente la hipótesis del auto-enmascaramiento, pero éste tema permanece irresuelto. (2) Dos de las características derivadas de las alas que aparentemente evolucionaron para suprimir el sonido del vuelo son los flecos del vexilo y la felpa dorsal de las alas de las lechuzas. Estas características ¿suprimen el ruido aerodinámico (sonido generado por el flujo de aire) o reducen en cambio el ruido estructural, tal como el ruido friccional de las plumas frotándose durante el vuelo? La hipótesis del ruido aerodinámico carece de apoyo empírico. Por el contrario, varias líneas de evidencia apoyan la hipótesis de que la felpa y el fleco reducen los sonidos friccionales, incluyendo: la posición anatómica del fleco y felpa, esta última mejor desarrollada en regiones del ala y cola propensos a frotación, y no tanto en áreas expuestas a flujo de aire ; la signatura acústica de frotación, que es de banda ancha e incluye ultrasonido, está presente en el vuelo de otras aves pero no en lechuzas; y la aparente relación entre la felpa y las bárbulas de fricción presentes en las remiges de otras aves. (3) ¿Evolucionó el vuelo silencioso en otros animales? Las características de las alas de las aves nocturnas (miembros nocturnos de Caprimulgiformes) sugieren que podrían haber evolucionado independientemente para volar de forma relativamente silenciosa, tal como ocurre en más de un gavilán diurno (Accipitriformes). Hipotetizamos que el vuelo de las aves es ruidoso porque las plumas alares están intrínsecamente predispuestas a frotarse y producir ruido friccional. La hipótesis sugiere una nueva perspectiva; en vez de considerar a las lechuzas como silenciosas, tal vez es que el vuelo de las aves es ruidoso. Esto implica que los murciélagos podrían representar un modelo ignorado de vuelo silencioso. El vuelo de las lechuzas podría no ser el mejor (y ciertamente no el único) modelo para la "bio-inspiración" del vuelo silencioso.Palabras clave: Acústica, Aeroacústica, sonidos inducidos por locomoción, Strigiformes, ala.

Subtle, pervasive genetic correlation between the sexes in the evolution of dimorphic hummingbird tail ornaments.

Male hummingbirds have repeatedly evolved sexually dimorphic tails that they use as ornaments during courtship. We examine how male ornament evolution is reflected in female morphology. Lande's two-step model of the evolution of dimorphism predicts that γ (the genetic correlation between the sexes) causes trait elaboration to first evolve quickly in both sexes, then dimorphism evolves more slowly. On the hummingbird phylogeny, tail length does not fit this two-step model; although hummingbirds repeatedly evolved ornamental, elongated tails, dimorphism evolves on the same phylogenetic branch as elongation, implying that γ quickly evolves to be low over phylogenetic timescales. Male "bee" hummingbirds have evolved diverse rectrix shapes that they use to produce sound. Female morphologies exhibit subtle, pervasive correlations with male morphology. No female-adaptive hypotheses explain these correlations, since females do not also make sounds with their tail. Subtle shape similarity has arisen through the genetic correlation with males, and is subject to intralocus sexual conflict. Intralocus sexual conflict may produce increased phenotypic variation of female ornaments. Other evolutionary constraints on tail morphology include a developmental correlation between neighboring tail-feathers, biasing tail elaboration to occur most often at the ends of the feather tract (rectrix 5 or 1) and not the middle.

Effect of Intensive Glycemic Control on Risk of Lower Extremity Amputation.

BACKGROUND: Diabetes mellitus is a major risk factor for peripheral arterial disease and lower extremity amputation (LEA). We evaluated the effects of intensive glucose control (IGC) on risk of LEA in patients with type 2 diabetes during a randomized-controlled multicenter trial. STUDY DESIGN: The Action to Control Cardiovascular Risk in Diabetes (ACCORD) trial randomized patients with type 2 diabetes to IGC (HbA1c target < 6.0%) or standard glycemic control (SGC; HbA1c target 7.0% to 7.9%). Using analysis of mean HbA1c, we examined relationships between glycemic control and incident/recurrent LEA during the clinical trial/follow-up. RESULTS: Mean post-randomization HbA1c over the course of the trial and post-trial follow-up was 7.3% ± 0.9% (6.8% ± 0.8% in the IGC arm, 7.7% ± 0.7% in the SGC arm). There were 124 participants who had at least 1 LEA during the study period; 73 were randomized to the SGC arm and 51 to the IGC arm (p = 0.049). Randomization to IGC was associated with decreased LEA rate (HR 0.69, 95% CI 0.483 to 0.987, p = 0.042). In multivariable models, mean HbA1c was a powerful predictor of LEA (HR 2.07 per 1% increase in HbA1c, 95% CI 1.67 to 2.57, p < 0.0001). Post-randomization mean HbA1c remained a strong predictor of LEA after controlling for other important covariates and competing risk of death (HR 1.94 per 1% increase in HbA1c, 95% CI 1.52 to 2.46, p < 0.0001). CONCLUSIONS: In patients with type 2 diabetes, IGC was associated with a reduction in the risk for LEA. After 3.7 years of IGC, there was an enduring protective effect against LEA. Improved glycemic control was a strong predictor of decreased risk for subsequent LEA. This study suggests that tight glycemic control, even over a short time period, has potential to reduce risk of limb loss.

Kinematic control of male Allen's hummingbird wing trill over a range of flight speeds.

Wing trills are pulsed sounds produced by modified wing feathers at one or more specific points in time during a wingbeat. Male Allen's hummingbirds (Selasphorus sasin) produce a sexually dimorphic 9 kHz wing trill in flight. Here, we investigated the kinematic basis for trill production. The wingtip velocity hypothesis posits that trill production is modulated by the airspeed of the wingtip at some point during the wingbeat, whereas the wing rotation hypothesis posits that trill production is instead modulated by wing rotation kinematics. To test these hypotheses, we flew six male Allen's hummingbirds in an open-jet wind tunnel at flight speeds of 0, 3, 6, 9, 12 and 14 m s-1, and recorded their flight with two 'acoustic cameras' placed below and behind, or below and lateral to the flying bird. The acoustic cameras are phased arrays of 40 microphones that used beamforming to spatially locate sound sources within a camera image. Trill sound pressure level (SPL) exhibited a U-shaped relationship with flight speed in all three camera positions. SPL was greatest perpendicular to the stroke plane. Acoustic camera videos suggest that the trill is produced during supination. The trill was up to 20 dB louder during maneuvers than it was during steady-state flight in the wind tunnel, across all airspeeds tested. These data provide partial support for the wing rotation hypothesis. Altered wing rotation kinematics could allow male Allen's hummingbirds to modulate trill production in social contexts such as courtship displays.

Quantification of free sugars, fructan, pungency and sweetness indices in onion populations by FT-MIR spectroscopy.

BACKGROUND: To facilitate faster phenotyping of onions (Allium cepa L.), Fourier-transform mid infrared (FT-MIR) spectroscopy with partial least squares (PLS) regression modelling was evaluated for the determination of pungency (pyruvate), sweetness (free sugars) and fructan in juice samples (n = 605) expressed from bulbs from breeding populations. RESULTS: Fourier-transform infrared (FTIR) spectra (range 1700-900 cm-1 ) were obtained from droplets (30 µL) of unprocessed juice. Goodness-of-fit (r2 ) and prediction errors (standard error of cross validation) for optimal PLS models were: soluble solids (0.997, 0.1 °Brix), pyruvate [0.825, 0.8 µmol g-1 fresh weight (FW)], fructan (0.98, 1.9 mg g-1 FW), glucose (0.941, 1.1 mg g-1 FW), fructose (0.967, 1.0 mg g-1 FW) and sucrose (0.919, 1.7 mg g-1 FW). FTIR models for industry sweetness indices based on glucose or sucrose equivalents were also developed. Because of its very low concentration (0.8-12 µmol g-1 FW) relative to other compounds, pyruvate was the weakest model developed. Fructan could be determined spectroscopically without the need for enzymatic digestion. CONCLUSIONS: All of the chemometric models developed are acceptable for screening purposes. Those for soluble solids, fructan and fructose are also suitable for routine analysis. FT-MIR can therefore be utilised for the simultaneous determination of pungency, sweetness and fructan in this crop. © 2018 Society of Chemical Industry.

Strategic Acoustic Control of a Hummingbird Courtship Dive.

Male hummingbirds court females with a high-speed dive in which they "sing" with their tail feathers. The male's choice of trajectory provides him strategic control over acoustic frequency and pressure levels heard by the female. Unlike related species, male Costa's hummingbirds (Calypte costae) choose to place their dives to the side of females. Here we show that this minimizes an audible Doppler curve in their dive sound, thereby depriving females of an acoustic indicator that would otherwise reveal male dive speed. Wind-tunnel experiments indicate that the sounds produced by their feathers are directional; thus, males should aim their tail toward females. High-speed video of dives reveal that males twist half of their tail vertically during the dive, which acoustic-camera video shows effectively aims this sound sideways, toward the female. Our results demonstrate that male animals can strategically modulate female perception of dynamic aspects of athletic motor displays, such as their speed.

Complex coevolution of wing, tail, and vocal sounds of courting male bee hummingbirds.

Phenotypic characters with a complex physical basis may have a correspondingly complex evolutionary history. Males in the "bee" hummingbird clade court females with sound from tail-feathers, which flutter during display dives. On a phylogeny of 35 species, flutter sound frequency evolves as a gradual, continuous character on most branches. But on at least six internal branches fall two types of major, saltational changes: mode of flutter changes, or the feather that is the sound source changes, causing frequency to jump from one discrete value to another. In addition to their tail "instruments," males also court females with sound from their syrinx and wing feathers, and may transfer or switch instruments over evolutionary time. In support of this, we found a negative phylogenetic correlation between presence of wing trills and singing. We hypothesize this transference occurs because wing trills and vocal songs serve similar functions and are thus redundant. There are also three independent origins of self-convergence of multiple signals, in which the same species produces both a vocal (sung) frequency sweep, and a highly similar nonvocal sound. Moreover, production of vocal, learned song has been lost repeatedly. Male bee hummingbirds court females with a diverse, coevolving array of acoustic traits.

Resonance frequencies of honeybee (Apis mellifera) wings.

During flight, insect wings bend and twist under the influence of aerodynamic and inertial forces. We tested whether wing resonance of honeybees (Apis mellifera) matches the wingbeat frequency, against the 'stiff element' hypothesis that the wing's first longitudinal mode exceeds the wingbeat frequency. Six bees were immobilized with their right wing pair outspread, and stimulated with a shaker while the normal modes were recorded with a scanning Doppler laser vibrometer. The lowest normal mode of the wings was the first longitudinal bending mode and, at 602±145 Hz, was greater than the wingbeat frequency of 234±13.9 Hz. Higher-order normal modes of the wing tended to incorporate nodal lines in the chordwise direction of the trailing edge, suggesting that their mode shape did not strongly resemble wing deformation during flapping flight. These results support the stiff element hypothesis for Apis mellifera.