RESUMO
How does agility evolve? This question is challenging because natural movement has many degrees of freedom and can be influenced by multiple traits. We used computer vision to record thousands of translations, rotations, and turns from more than 200 hummingbirds from 25 species, revealing that distinct performance metrics are correlated and that species diverge in their maneuvering style. Our analysis demonstrates that the enhanced maneuverability of larger species is explained by their proportionately greater muscle capacity and lower wing loading. Fast acceleration maneuvers evolve by recruiting changes in muscle capacity, whereas fast rotations and sharp turns evolve by recruiting changes in wing morphology. Both species and individuals use turns that play to their strengths. These results demonstrate how both skill and biomechanical traits shape maneuvering behavior.
Assuntos
Aves/anatomia & histologia , Aves/fisiologia , Voo Animal/fisiologia , Músculo Esquelético/anatomia & histologia , Músculo Esquelético/fisiologia , Asas de Animais/anatomia & histologia , Asas de Animais/fisiologia , Aceleração , Animais , Evolução Biológica , Aves/classificação , Filogenia , Rotação , América do SulRESUMO
The tempo of species diversification in large clades can reveal fundamental evolutionary mechanisms that operate on large temporal and spatial scales. Hummingbirds have radiated into a diverse assemblage of specialized nectarivores comprising 338 species, but their evolutionary history has not, until now, been comprehensively explored. We studied hummingbird diversification by estimating a time-calibrated phylogeny for 284 hummingbird species, demonstrating that hummingbirds invaded South America by â¼22 million years ago, and subsequently diversified into nine principal clades (see [5-7]). Using ancestral state reconstruction and diversification analyses, we (1) estimate the age of the crown-group hummingbird assemblage, (2) investigate the timing and patterns of lineage accumulation for hummingbirds overall and regionally, and (3) evaluate the role of Andean uplift in hummingbird speciation. Detailed analyses reveal disparate clade-specific processes that allowed for ongoing species diversification. One factor was significant variation among clades in diversification rates. For example, the nine principal clades of hummingbirds exhibit â¼15-fold variation in net diversification rates, with evidence for accelerated speciation of a clade that includes the Bee, Emerald, and Mountain Gem groups of hummingbirds. A second factor was colonization of key geographic regions, which opened up new ecological niches. For example, some clades diversified in the context of the uplift of the Andes Mountains, whereas others were affected by the formation of the Panamanian land bridge. Finally, although species accumulation is slowing in all groups of hummingbirds, several major clades maintain rapid rates of diversification on par with classical examples of rapid adaptive radiation.
Assuntos
Distribuição Animal , Biodiversidade , Aves/genética , Evolução Molecular , Especiação Genética , Filogenia , Animais , Sequência de Bases , Teorema de Bayes , Geografia , Modelos Genéticos , Dados de Sequência Molecular , América do Norte , Análise de Sequência de DNA , América do Sul , Especificidade da EspécieRESUMO
Hummingbirds, with their impressive flight ability and competitive aerial contests, make ideal candidates for applying a mechanistic approach to studying community structure. Because flight costs are influenced by abiotic factors that change systematically with altitude, elevational gradients provide natural experiments for hummingbird flight ecology. Prior attempts relied on wing disc loading (WDL) as a morphological surrogate for flight performance, but recent analyses indicate this variable does not influence either territorial behavior or competitive ability. Aerodynamic power, by contrast, can be derived from direct measurements of performance and, like WDL, declines across elevations. Here, I demonstrate for a diverse community of Andean hummingbirds that burst aerodynamic power is associated with territorial behavior. Along a second elevational gradient in Colorado, I tested for correlated changes in aerodynamic power and competitive ability in two territorial hummingbirds. This behavioral analysis revealed that short-winged Selasphorus rufus males are dominant over long-winged Selasphorus platycercus males at low elevations but that the roles are reversed at higher elevations. Several lines of evidence support the hypothesis that the burst rather than sustained aerodynamic performance mediates competitive ability at high elevation. A minimum value for burst power may be required for successful competition, but other maneuverability features gain importance when all competitors have sufficient muscle power, as occurs at low elevations.
Assuntos
Aves/fisiologia , Comportamento Competitivo , Meio Ambiente , Voo Animal/fisiologia , Animais , Aves/anatomia & histologia , Colorado , Peru , Predomínio Social , Territorialidade , Asas de Animais/anatomia & histologia , Asas de Animais/fisiologiaRESUMO
Hovering flight is one of the most energetically demanding forms of animal locomotion. Despite the cost, hummingbirds regularly hover at high elevations, where flight is doubly challenging because of reduced air density and oxygen availability. We performed three laboratory experiments to examine how air density and oxygen partial pressure influence wingbeat kinematics. In the first study, we experimentally lowered air density but maintained constant oxygen partial pressure. Under these hypodense but normoxic conditions, hummingbirds increased stroke amplitude substantially and increased wingbeat frequency slightly. In the second experiment, we maintained constant air density but decreased oxygen partial pressure. Under these normodense but hypoxic conditions, hummingbirds did not alter stroke amplitude but instead reduced wingbeat frequency until they could no longer generate enough vertical force to offset body weight. In a final combined experiment, we decreased air density but increased oxygen availability, and found that the wingbeat kinematics were unaffected by supplemental oxygen. We also studied hovering and maximally loaded flight performance for 43 hummingbird species distributed along a natural elevational gradient in Peru. During free hovering flight, hummingbirds showed increased stroke amplitude interspecifically at higher elevations, mirroring the intra-individual responses in our first laboratory experiment. During loaded flight, hummingbirds increased both wingbeat frequency and wing stroke amplitude by 19% relative to free-flight values at any given elevation. We conclude that modulation of wing stroke amplitude is a major compensatory mechanism for flight in hypodense or hypobaric environments. By contrast, increases in wingbeat frequency impose substantial metabolic demands, are only elicited transiently and anaerobically, and cannot be used to generate additional sustained lift at high elevations.