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Shawn R. Craik, Amélie Doucet, Mathieu Manuel, Chloé Roy, Taylor M. Brown, Emilie J. Knighton, Dave Shutler, Randy F. Lauff, Danielle Ethier, Amy-Lee Kouwenberg, Philip D. Taylor
KEYWORDS: Aegolius acadicus, body condition, breeding abundance, capture date, Density-dependent effects, irruptive migration, Northern Saw-whet Owl, Stopover duration, Aegolius acadicus, condition physique, abondance de reproducteurs, date de capture, effets dépendant de la densité, migration irruptive, durée de la halte migratoire
Bird species that undertake irruptive migrations are good candidates for assessing density-dependent effects on stopover ecology because the number of birds using stopover sites varies considerably from year to year. Using morphometric data from a 9-year banding program and radiotelemetry (n = 25 females), we found that increases in the annual density of Northern Saw-whet Owls (NSWO; Aegolius acadicus) encountered at a fall stopover site in Nova Scotia, Canada, were not linked to female body condition or minimum stopover duration. Rather, most NSWO spent no more than 1 or 2 full days at the stopover site following radio-tagging and during their return visits to the site. Body condition indices were highest for NSWO captured near the end of the migration monitoring season, possibly reflecting birds that had recently established wintering ranges and were elevating energy stores. We used breeding abundance indices derived from Birds Canada's Atlantic Nocturnal Owl Survey to help test the hypothesis that irruptive fall migrations in NSWO were driven by elevated breeding productivity and dispersal of immature birds (breeding success hypothesis). Indeed, irruptive fall migrations were characterized by elevated densities of immatures, but not adults, and mean breeding abundance indices for the Maritime provinces during years with irruptive migrations were higher than those for non-irruptive years. We hypothesize that prey abundance during years with irruptive migrations was sufficient to enable high breeding densities and minimize effects of elevated fall densities of NSWO on rates of mass gain and stopover duration.
LAY SUMMARY
Elevated migrant densities can affect a bird's rate of mass gain, and individuals in poor condition may need to extend duration of their stopovers to acquire energy to resume migration.
Using a banding dataset and radiotelemetry, we assessed whether densities of Northern Saw-whet Owls (Aegolius acadicus; NSWO) encountered at a fall stopover site in Nova Scotia, Canada, were associated with owl body condition and stopover duration.
Annual densities of NSWO detected at the stopover site varied across the 9-year study period; however, female body condition and stopover duration were not linked to owl densities.
Irruptive migrations consisted of elevated densities of immature, but not adult, NSWO, and mean breeding abundance indices for years with irruptive migrations were higher than those for non-irruptive years, consistent with the hypothesis that irruptive movements can be driven by high breeding productivity in this species.
We hypothesize that prey availability during years with irruptive fall migrations was sufficient to enable elevated breeding abundance of NSWO and reduce effects of increased fall densities on rates of mass gain and stopover duration.
Les espèces d'oiseaux qui entreprennent des migrations irruptives sont de bons candidats pour évaluer les effets dépendant de la densité sur l'écologie de la halte migratoire car le nombre d'oiseaux utilisant les sites de halte migratoire varie considérablement d'une année à l'autre. En utilisant les données morphométriques provenant d'un programme de baguage s'étendant sur 9 ans et la radiotélémétrie (n = 25 femelles), nous avons trouvé que les hausses de densité annuelle d'Aegolius acadicus (NSWO) rencontrées sur un site de halte migratoire automnale en Nouvelle-Écosse, au Canada, n'étaient pas liées à la condition physique des femelles ni à une durée minimale de halte migratoire. Au contraire, la plupart des NSWO n'ont pas passé plus d'un ou deux jours complets au site de halte migratoire après avoir été munis d'un émetteur et lors de leurs visites de retour sur le site. Les indices de condition physique étaient plus élevés pour les NSWO capturés vers la fin de la saison de suivi de la migration, lesquels pourraient être des oiseaux ayant récemment établi leur aire d'hivernage et qui étaient en train d'accumuler des réserves d'énergie. Nous avons utilisé des indices de l'abondance de reproducteurs dérivés de l'Inventaire des Strigidés Nocturnes de l'Atlantique d'Oiseaux Canada pour vérifier l'hypothèse que les migrations automnales irruptives chez NSWO étaient motivées par une productivité reproductive élevée et la dispersion d'oiseaux immatures (hypothèse du succès de la reproduction). En effet, les migrations automnales irruptives étaient caractérisées par des densités élevées d'immatures, mais pas d'adultes, et les valeurs moyennes des indices de l'abondance de reproducteurs pour les provinces maritimes pendant les années de migrations irruptives étaient plus élevées que celles des années sans irruption. Nous émettons l'hypothèse que l'abondance des proies durant les années avec des migrations irruptives était suffisante pour permettre des densités de reproduction élevées et limiter les effets des densités automnales élevées de NSWO sur les taux d'accroissement de la masse et la durée de la halte migratoire.
In high-latitude species with high dispersal ability, such as long-distance migratory birds, populations are often assumed to exhibit little genetic structure due to high gene flow or recent postglacial expansion. We sequenced over 120 low-coverage whole genomes from across the breeding range of a long-distance migratory bird, the Veery (Catharus fuscescens), revealing strong evidence for isolation by distance. Additionally, we found distinct genetic structure between boreal, western montane U.S., and southern Appalachian sampling regions. We suggest that population genetic structure in this highly migratory species is detectable with the high resolution afforded by whole-genomic data because, similar to many migratory birds, the Veery exhibits high breeding-site fidelity, which likely limits gene flow. Resolution of isolation by distance across the breeding range was sufficient to assign likely breeding origins of individuals sampled in this species' poorly understood South American nonbreeding range, demonstrating the potential to assess migratory connectivity in this species using genomic data. As the Veery's breeding range extends across both historically glaciated and unglaciated regions in North America, we also evaluated whether contemporary patterns of structure and genetic diversity are consistent with historical population isolation in glacial refugia. We found that patterns of genetic diversity did not support southern montane regions (southern Appalachians or western U.S. mountains) as glacial refugia. Overall, our findings suggest that isolation by distance yields subtle associations between genetic structure and geography across the breeding range of this highly vagile species even in the absence of obvious historical vicariance or contemporary barriers to dispersal.
LAY SUMMARY
Describing how populations in a species differ genetically is important for understanding that species' evolutionary history.
Migratory birds have high dispersal abilities, potentially reducing genetic structure. However, many migratory birds return to the same breeding site year after year, which could reduce gene flow between populations.
We sequenced >120 genomes to detect population genetic differentiation in a common songbird, the Veery (Catharus fuscescens).
We found that genetic similarity between samples decreased with increasing geographic distance (i.e., isolation by distance) and that populations in southern Appalachia were distinct from samples in the rest of the breeding range.
We did not detect differences in genetic diversity patterns between populations, contrary to predictions about putative glacial refugia.
We assigned likely geographic breeding region to birds sampled in the poorly understood wintering range.
Despite the Veery's long-distance migration and high dispersal ability, isolation by distance produces subtle but detectable population structure across its breeding range.
En especies de latitudes altas con elevada capacidad de dispersión, como las aves migratorias de larga distancia, a menudo se asume que las poblaciones exhiben poca estructura genética debido al alto flujo génico o a la reciente expansión post-glaciar. Secuenciamos más de 120 genomas completos de baja cobertura en todo el rango de cría de un ave migratoria de larga distancia, Catharus fuscescens, revelando fuertes evidencias de aislamiento por distancia. Además, encontramos una estructura genética distintiva entre las regiones de muestreo boreales, las montañas del oeste de EEUU y los Montes Apalaches del sur. Sugerimos que la estructura genética poblacional en esta especie altamente migratoria es detectable por la alta resolución que proporcionan los datos genómicos completos porque, al igual que muchas aves migratorias, C. fuscescens muestra una alta fidelidad al sitio de cría, lo que probablemente limita el flujo génico. La resolución del aislamiento por distancia en todo el rango de cría fue suficiente para asignar los probables orígenes de cría de individuos muestreados en el rango no reproductivo -y poco conocido- de esta especie en Sudamérica, demostrando el potencial para evaluar la conectividad migratoria en esta especie utilizando datos genómicos. Dado que el rango de cría de C. fuscescens se extiende por regiones históricamente con y sin glaciares en América del Norte, también evaluamos si los patrones contemporáneos de estructura y diversidad genética son consistentes con el aislamiento poblacional histórico en refugios glaciares. Encontramos que los patrones de diversidad genética no respaldaban a las regiones montañosas del sur (Montes Apalaches del sur o montañas del oeste de EEUU) como refugios glaciares. En general, nuestros hallazgos sugieren que el aislamiento por distancia produce asociaciones sutiles entre la estructura genética y la geografía en todo el rango de cría de esta especie altamente móvil, incluso en ausencia de vicarianzas históricas obvias o de barreras contemporáneas para la dispersión.
Previous work has demonstrated that there is extensive variation in the songs of White-crowned Sparrow (Zonotrichia leucophrys) throughout the species range, including between neighboring (and genetically distinct) subspecies Z. l. nuttalli and Z. l. pugetensis. Using a machine learning approach to bioacoustic analysis, we demonstrate that variation in song is correlated with year of recording (representing cultural drift), geographic distance, and climatic differences, but the response is subspecies- and season-specific. Automated machine learning methods of bird song annotation can process large datasets more efficiently, allowing us to examine 1,913 recordings across ∼60 years. We utilize a recently published artificial neural network to automatically annotate White-crowned Sparrow vocalizations. By analyzing differences in syllable usage and composition, we recapitulate the known pattern where Z. l. nuttalli and Z. l. pugetensis have significantly different songs. Our results are consistent with the interpretation that these differences are caused by the changes in characteristics of syllables in the White-crowned Sparrow repertoire. This supports the hypothesis that the evolution of vocalization behavior is affected by the environment, in addition to population structure.
LAY SUMMARY
Birdsong is an important behavior because it is critical to bird communication and reproduction.
White-crowned Sparrows in western North America are known to use different songs along their range, but it is unknown how those songs vary through space and time.
We used machine learning to analyze these songs and found that populations of White-crowned Sparrows can be differentiated based on their songs.
Geographic factors exert a greater influence on song evolution in migratory populations, while climate factors are important across all populations.
Songs in this species also change through time irrespective of other changes, meaning they are impacted by cultural evolution.
Trabajos anteriores han demostrado que existe una extensa variación en los cantos de Zonotrichia leucophrys en todo el rango de la especie, incluida la variación entre las subespecies vecinas (y genéticamente distintas) Z. l. nuttalli y Z. l. pugetensis. Utilizando un enfoque de aprendizaje automático para el análisis bioacústico, demostramos que la variación en el canto está correlacionada con el año de grabación (que representa la deriva cultural), la distancia geográfica y las diferencias climáticas, pero la respuesta es específica de la subespecie y la temporada. Los métodos automatizados de aprendizaje automático para analizar el canto de las aves pueden procesar sets de datos grandes de manera más eficiente, lo que nos permitió examinar 1,913 grabaciones realizadas a lo largo de ∼60 años. Utilizamos una red neuronal artificial recientemente publicada para analizar automáticamente las vocalizaciones de Z. leucophrys. Al analizar las diferencias en el uso y la composición de las sílabas, recapitulamos el patrón conocido en el que Z. l. nuttalli y Z. l. pugetensis tienen cantos significativamente diferentes. Nuestros resultados son consistentes con la interpretación de que estas diferencias son causadas por los cambios en las características de las sílabas en el repertorio de Z. leucophrys. Esto respalda la hipótesis de que la evolución del comportamiento de vocalización se ve afectada por el ambiente, además del efecto de la estructura poblacional.
We assessed correlations between speciation rate and bill morphology in an ecologically diverse and continental-scale songbird radiation, the tanagers (Thraupidae). Our analyses showed that bill size, shape, and their evolutionary rates are not correlated with speciation rate. However, we did find evidence that each axis of variation in bill morphology diversifies at speciation events interspersed with periods of gradual evolution, consistent with a punctuated equilibrium model of character change. To determine correlations, we incorporated a time-calibrated molecular phylogeny and high-resolution three-dimensional surface scans of bill structure from museum study skins. Overall, our findings suggest that bill size and shape diversify in association with the speciation process, but not through any effect on the rate of speciation. Previous studies have shown other traits, such as song, ecological niche, and plumage do influence speciation and, ultimately, species richness in tanagers.
LAY SUMMARY
Bill morphology is widely considered to play a role in avian speciation and diversification as a result of ecological specialization
Previous studies on the relationship between bill morphology and speciation have mostly focused on closely related species often in island settings
We quantify correlations between speciation rate and bill morphology in a diverse and broadly distributed songbird radiation, the tanagers
We find that bill size, shape, and their evolutionary rates are not correlated with speciation rate
We did find support for mixed punctuated and gradual evolution of bill morphology
Our results suggest that bill size and shape diversify in association with the speciation process, but not through any effect on the rate of speciation in the tanagers
Evaluamos las correlaciones entre la tasa de especiación y la morfología del pico (tamaño, forma y tasa evolutiva) en una radiación diversa y ampliamente distribuida de aves canoras, los tangaras (Thraupidae). Nuestros análisis mostraron que el tamaño, la forma y las tasas evolutivas del pico no están correlacionadas con la tasa de especiación. Sin embargo, encontramos evidencia de que cada eje de variación en la morfología del pico se diversifica en eventos de especiación intercalados con períodos de evolución gradual, en consonancia con un modelo de equilibrio puntuado de cambio de caracteres. Para determinar las correlaciones, incorporamos una filogenia molecular calibrada en el tiempo y escaneos de superficie de alta resolución en 3D de la estructura del pico a partir de pieles de estudio de museos. En general, nuestros hallazgos sugieren que el tamaño y la forma del pico se diversifican en asociación con el proceso de especiación, pero no a través de ningún efecto en la tasa de especiación. Estudios previos han demostrado que otros rasgos, como el canto, el nicho ecológico y el plumaje, sí influyen en la especiación y, en última instancia, en la riqueza de especies en los tangaras.
Most of our knowledge of secondary hole-nesting birds comes from populations breeding in human-provided nestboxes, yet these might differ from natural cavities in a number of parameters, including internal dimensions or microclimate, leading to differences in reproductive ecology. Here, we demonstrate differences in plumage coloration, an important visual signal of individual quality, in Blue Tit (Cyanistes caeruleus) and Great Tit (Parus major) nestlings raised in natural cavities and nestboxes. For this study, we collected feather samples over two breeding seasons and applied reflectance spectrophotometry to calculate color metrics. Carotenoid chroma was higher in nestbox-reared Blue Tits, whereas brightness was higher in nestbox-reared Great Tits (with a similar tendency for Blue Tits). The latter result might be explained by the better nutritional condition of Great Tit nestlings raised in nestboxes. Furthermore, we found no evidence for preference of adults expressing more elaborate plumage towards a specific cavity type in either species. Consequently, we assume that differences in nestling plumage reported here are driven by rearing conditions (nestboxes vs. natural cavities) and not by differences in plumage-based parental quality. Our study adds to the growing body of evidence confirming that anthropogenic environmental modifiers, such as nestboxes, might influence avian physiology and the resulting phenotype.
LAY SUMMARY
Natural cavities and nestboxes differ in many ways, including dimensions and microclimatic conditions, which can have consequences for predation risk and avian reproductive success.
Currently, knowledge on plumage color of hole-nesters comes solely from studies on populations breeding in nestboxes or aviaries. So far, no study tested whether such results hold for birds breeding in natural cavities. To fill this gap, we examined the impact of rearing cavity type—natural cavities vs. artificial cavities (nestboxes) on carotenoid-based coloration of Blue Tit (Cyanistes caeruleus) and Great Tit (Parus major) nestlings.
We found that BlueTit nestlings expressed plumage with more saturated color in nestboxes, whereas in GreatTits nestlings raised in nestboxes expressed brighter plumage.
Our results add to the growing body of evidence that differences between natural cavities and nestboxes can bear consequences for nestling physiology and phenotype.
La plupart de nos connaissances sur les oiseaux cavicoles secondaires proviennent de populations se reproduisant dans des nichoirs artificiels, mais ceux-ci peuvent différer des cavités naturelles d'un certain nombre de paramètres, incluant les dimensions internes ou le microclimat, ce qui entraîne des différences dans l'écologie de la reproduction. Dans cette étude, nous démontrons qu'il existe des différences dans la coloration du plumage, un signal visuel important de la qualité individuelle, chez les oisillons de Cyanistes caeruleus et Parus major élevés dans des cavités naturelles et dans des nichoirs. Nous avons collecté des échantillons de plumes pendant deux saisons de reproduction et nous avons appliqué la spectrophotométrie de réflectance pour calculer les paramètres de la couleur. La saturation des couleurs des caroténoïdes était plus élevée chez les oisillons de C. caeruleus élevés dans des nichoirs, alors que la brillance était plus élevée chez les oisillons de P. major élevés dans des nichoirs (avec une tendance similaire pour C. caeruleus). Ce dernier résultat pourrait s'expliquer par le meilleur état nutritionnel des oisillons de P. major élevés dans des nichoirs. En outre, nous n'avons trouvé aucune preuve de la préférence des adultes au plumage plus élaboré pour un type de cavité spécifique chez l'une ou l'autre de ces espèces. Par conséquent, nous supposons que les différences de plumage chez les oisillons sont dues aux conditions d'élevage (nichoirs vs cavités naturelles) et non à des différences de qualité parentale basées sur le plumage. Notre étude s'ajoute au nombre croissant de preuves confirmant que les modificateurs environnementaux anthropogéniques, tels que les nichoirs, peuvent influencer la physiologie aviaire et le phénotype qui en résulte.
Molt is an essential life-history event in birds and many mammals, as maintenance of feathers and fur is critical for survival. Despite this molt remains an understudied life-history event. Non-standard statistical techniques are required to estimate the phenology of molt from observations of plumage or pelage state, and existing molt phenology models have strict sampling requirements that can be difficult to meet under real-world conditions. We present an extended modelling framework that can accommodate features of real-world molt datasets, such as re-encounters of individuals, misclassified molt states, and/or molt state-dependent sampling bias. We demonstrate that such features can lead to biased inferences when using existing molt phenology models, and show that our model extensions can improve inferences about molt phenology under a wide range of sampling conditions. We hope that our novel modelling framework removes barriers for modelling molt phenology data from real-world datasets and thereby further facilitates the uptake of appropriate statistical methods for such data. Although we focus on molt, the modelling framework is applicable to other phenological processes that can be recorded using either ordered categories or approximately linear progress scores.
LAY SUMMARY
Molt is an essential but understudied life-history event in birds and many mammals.
The analysis of molt data requires non-standard statistical techniques and existing molt phenology models have strict sampling requirements that can be difficult to meet.
We present new statistical models for molt timing and duration.
We demonstrate that these models improve inferences about molt duration and timing under a wide range of realistic sampling conditions.
La muda es un evento esencial en la historia de vida de las aves y de muchos mamíferos, ya que el mantenimiento de las plumas y del pelaje es crucial para la supervivencia. A pesar de esto, la muda sigue siendo un evento de la historia de vida poco estudiado. Se requieren técnicas estadísticas no convencionales para estimar la fenología de la muda a partir de observaciones del estado del plumaje o pelaje, y los modelos de fenología de la muda existentes tienen requisitos estrictos de muestreo que pueden ser difíciles de cumplir en condiciones del mundo real. Presentamos un marco de modelado extendido que puede adaptarse a las características de los sets de datos de la muda del mundo real, como reencuentros de individuos, estados de la muda mal clasificados y/o sesgo de muestreo dependiente del estado de la muda. Demostramos que tales características pueden llevar a inferencias sesgadas al usar modelos existentes de fenología de la muda, y presentamos cómo nuestras extensiones del modelo pueden mejorar las inferencias sobre la fenología de la muda en una amplia gama de condiciones de muestreo. Esperamos que nuestro nuevo marco de modelado elimine las barreras existentes para modelar los datos de fenología de la muda de los sets de datos del mundo real y, de esta manera, facilite aún más la adopción de métodos estadísticos apropiados para tales datos. Aunque nos centramos en la muda, el marco de modelado es aplicable a otros procesos fenológicos que pueden registrarse mediante categorías ordenadas o clasificaciones de progreso aproximadamente lineales.
KEYWORDS: birds, clutch size, islands, island syndrome, latitude, life history, phylogenetic, seabirds, Aves, aves marinas, filogenético, historia de vida, islas, latitud, síndrome de la isla, tamaño de nidada
The “island syndrome” refers to similarity in the biology of island organisms, but its generality is questionable, as the scope of species and traits examined are often limited. Here, I show that birds breeding exclusively on islands (breeding island endemics) evolved smaller clutches, using a dataset of 4,530 bird species. Using an inclusive definition of a breeding island endemic, which also encompasses migratory species and seabirds, I examine the evolution of clutch sizes in island breeding species using phylogenetic generalized linear models. Across disparate phylogenetic hypotheses, and after accounting for biological and geographical co-variables, I show that breeding island endemic landbirds (470 species) evolved smaller clutch sizes than continental breeding species (3,818 species). I show that the evolution of clutch size follows the expectations of the island syndrome, as among breeding island endemic landbirds there is a positive relationship between clutch size and breeding range area. Finally, I reinforce the view that the island syndrome is a general pattern in birds, spanning diverse phylogenetic and ecological groups, by showing that in a seabird-only dataset (242 species), breeding island endemic seabirds show evolution of smaller clutch sizes. In a model of the full dataset of both landbirds and seabirds (4,530 species) there was no evidence of an interaction of being a seabird with breeding island endemicity, showing that seabirds and landbirds respond in the same way. This study, using more than 40% of all bird species, provides the first evidence of a general evolutionary response in a life history trait, clearly showing the island syndrome as a general evolutionary tendency associated with island environments.
LAY SUMMARY
The island syndrome is a phenomenon of similarity in morphology, behavior, and life history between island organisms.
Even among birds, its extent is not fully studied; it is not clear which traits (particularly behavioral and life history traits) are involved, and how generally it applies across the whole diversity of birds (more than 10,000 species).
Using a dataset of 4,530 species with established molecular phylogenies, I examined whether birds that nest exclusively on islands consistently differ from continental-breeding species in terms of a key life history trait: clutch size.
I found that breeding island endemic landbirds have smaller clutches. Among island breeding endemic landbirds, breeding range area was positively correlated with clutch size, as expected in the island syndrome. Seabirds show the same evolutionary response in clutch size as landbirds.
This study shows that a life history trait is part of the island syndrome across a representative sample of the avian phylogeny, and shows that seabirds also exhibit the island syndrome.
El “síndrome de la isla” se refiere a la similitud en la biología de los organismos de las islas, pero su generalidad es cuestionable, ya que la amplitud de especies y rasgos examinados a menudo es limitada. Aquí, muestro que las aves que se reproducen exclusivamente en islas (reproductoras endémicas de islas) evolucionaron nidadas más pequeñas, utilizando un set de datos de 4.530 especies de aves. Utilizando una definición inclusiva de reproductoras endémicas de islas, que también abarca especies migratorias y aves marinas, examino la evolución de los tamaños de nidada en especies reproductoras de islas mediante modelos lineales generalizados filogenéticos. A través de diversas hipótesis filogenéticas, y después de tener en cuenta las covariables biológicas y geográficas, demuestro que las aves terrestres reproductoras endémicas de islas (470 especies) evolucionaron tamaños de nidada más pequeños que las especies continentales (3.818 especies). Demuestro que la evolución del tamaño de la nidada sigue las expectativas del síndrome de la isla, ya que entre las aves terrestres reproductoras endémicas de islas hay una relación positiva entre el tamaño de la nidada y el área de reproducción. Finalmente, refuerzo la idea de que el síndrome de la isla es un patrón general en las aves, abarcando diversos grupos filogenéticos y ecológicos, al mostrar que en un set de datos exclusivo de aves marinas (242 especies), las aves marinas reproductoras endémicas de islas muestran la evolución de tamaños de nidada más pequeños. En un modelo del set completo de datos de aves terrestres y marinas (4.530 especies), no hubo evidencia de una interacción entre ser un ave marina y ser un ave reproductora endémica de islas, mostrando que las aves marinas y las aves terrestres responden de la misma manera. Este estudio, que utiliza más del 40% de todas las especies de aves, proporciona la primera evidencia de una respuesta evolutiva general en un rasgo de la historia de vida, mostrando claramente el síndrome de la isla como una tendencia evolutiva general asociada con ambientes isleños.
Many animals rely on photoperiodic and non-photoperiodic environmental cues to gather information and appropriately time life-history stages across the annual cycle, such as reproduction, molt, and migration. Here, we experimentally demonstrate that the reproductive physiology, but not migratory behavior, of captive Pine Siskins (Spinus pinus) responds to both food and social cues during the spring migratory-breeding period. Pine Siskins are a nomadic finch with a highly flexible breeding schedule and, in the spring, free-living Pine Siskins can wander large geographic areas and opportunistically breed. To understand the importance of non-photoperiodic cues to the migratory-breeding transition, we maintained individually housed birds on either a standard or enriched diet in the presence of group-housed heterospecifics or conspecifics experiencing either the standard or enriched diet type. We measured body condition and reproductive development of all Pine Siskins and, among individually housed Pine Siskins, quantified nocturnal migratory restlessness. In group-housed birds, the enriched diet caused increases in body condition and, among females, promoted reproductive development. Among individually housed birds, female reproductive development differed between treatment groups, whereas male reproductive development did not. Specifically, individually housed females showed greater reproductive development when presented with conspecifics compared to heterospecifics. The highest rate of female reproductive development, however, was observed among individually housed females provided the enriched diet and maintained with group-housed conspecifics on an enriched diet. Changes in nocturnal migratory restlessness did not vary by treatment group or sex. By manipulating both the physical and social environment, this study demonstrates how multiple environmental cues can affect the timing of transitions between life-history stages with differential responses between sexes and between migratory and reproductive systems.
LAY SUMMARY
Many animals inform decisions associated with survival and reproduction using a multitude of environmental cues, including photoperiodic and non-photoperiodic cues.
How multiple environmental cues combine to influence animal decision-making is not well understood.
We experimentally tested how different combinations of non-photoperiodic cues, including social cues, influenced the spring migratory behavior and reproductive physiology of Pine Siskins.
Pine Siskins are a nomadic finch with highly flexible breeding and migratory behavior that are known to be modulated by non-photoperiodic cues.
Results demonstrate that the presence of conspecifics—but not heterospecifics that are ecologically relevant—and direct access to preferred diet items promote female reproductive development.
Male reproductive development and nocturnal migratory restlessness did not differ across experimental treatments.
Our results highlight sex-specific differences in environmental cue use and demonstrate how animals can use multiple types of environmental cues to differentially inform decisions associated with the timing of annual cycle events.
Muchos animales dependen de señales ambientales fotoperiódicas y no fotoperiódicas para recopilar información y sincronizar adecuadamente las etapas del ciclo anual de su historia de vida, como la reproducción, la muda y la migración. Aquí, demostramos experimentalmente que la fisiología reproductiva, pero no el comportamiento migratorio, de individuos de Spinus pinus en cautiverio responden tanto a señales alimenticias como sociales durante el período migratorio-reproductivo de primavera. S. pinus es un ave nómada con un esquema reproductivo altamente flexible y, en primavera, los individuos que viven en libertad pueden deambular por grandes áreas geográficas y reproducirse de modo oportunista. Para entender la importancia de las señales no fotoperiódicas en la transición migratoria-reproductiva, mantuvimos aves alojadas individualmente ya sea con una dieta estándar o una enriquecida, en presencia de individuos heteroespecíficos o conspecíficos alojados en grupo, que recibieron ya sea el tipo de dieta estándar o la enriquecida. Medimos la condición corporal y el desarrollo reproductivo de todos los individuos de S. pinus y, entre los individuos alojados individualmente, cuantificamos la inquietud migratoria nocturna. En las aves alojadas en grupo, la dieta enriquecida causó aumentos en la condición corporal y, entre las hembras, promovió el desarrollo reproductivo. Entre las aves alojadas individualmente, el desarrollo reproductivo de las hembras difirió entre los grupos de tratamiento, mientras que el desarrollo reproductivo de los machos no lo hizo. En particular, las hembras alojadas individualmente mostraron un mayor desarrollo reproductivo cuando se presentaban con conspecíficos en comparación con heteroespecíficos. La tasa más alta de desarrollo reproductivo en las hembras, sin embargo, se observó entre las hembras alojadas individualmente que recibieron la dieta enriquecida y que se mantuvieron con conspecíficos alojados en grupo con la dieta enriquecida. Los cambios en la inquietud migratoria nocturna no variaron según el grupo de tratamiento o el sexo. Al manipular tanto el entorno físico como social, este estudio demuestra cómo múltiples señales ambientales pueden afectar el momento de las transiciones entre las etapas de la historia de vida, con respuestas diferenciales entre los sexos y entre los sistemas migratorios y reproductivos.
The passerine superfamily Certhioidea lacks a complete phylogeny despite decades of recognition as a clade and extensive systematic work within all its constituent families. Here, we inferred a near-complete species-level phylogeny of Certhioidea from a molecular supermatrix, including the first comprehensive sampling of the wrens (Troglodytidae), and used this phylogeny to infer its biogeographic and diversification histories. We also inferred an expanded phylogeny including nearly 100 putative phylospecies previously documented in the literature, and we found that including this diversity had notable impacts on the inferred evolutionary history of Certhioidea. This phylospecies-level tree documented a few instances of species paraphyly, some previously described in the literature and some novel. We found that Certhioidea originated largely in Eurasia and dispersed into North America five times in the last 20 million years, including at the origin of the “New World certhioids,” wrens and gnatcatchers, a clade herein named Orthourae. After this initial dispersal event, both wrens and gnatcatchers diversified extensively across the hemisphere, with both lineages repeatedly crossing between continents. However, we detected no notable impact of the formation of the Isthmus of Panama on the frequency of dispersal events between North and South America. The inclusion of phylospecies altered this biogeographic inference in some portions of the tree but overall was largely consistent. With species-level sampling, we found that diversification rates within Certhioidea were largely constant through time with a detectable deceleration toward the present. By contrast, phylospecies-level sampling recovered a different diversification history with a significant rate increase at the crown node of Orthourae after dispersing into the Americas and increased speciation rates particularly within the genera Polioptila and Henicorhina. This largely resolved phylogeny for Certhioidea has yielded important insights into the evolutionary history of this group and provides a framework for future comparative work on this fascinating clade.
LAY SUMMARY
Wrens are one of the most diverse songbird groups in the Americas, but we currently lack a complete understanding of their evolutionary history in relation to their relatives: the gnatcatchers, nuthatches, tree creepers, and Wallcreeper.
We addressed this gap in knowledge by combining existing and new sequence data to estimate a near-complete phylogenetic tree for the wrens and their allies that is largely consistent with previous studies, but not well resolved for a subgroup of the wrens.
This tree informed us that this group originated in Eurasia and has dispersed several times to Africa and North America in the last 20 million years.
By including nearly 100 phylospecies, we demonstrated that the potential underestimation of the diversity in this group may alter our understanding of this group's evolutionary history across space and time.
La superfamilia de aves paseriformes Certhioidea carece de una filogenia completa, a pesar de décadas de reconocimiento como un clado y de extenso trabajo sistemático dentro de todas sus familias constituyentes. Aquí, inferimos una filogenia casi completa a nivel de especie de Certhioidea a partir de una supermatriz molecular, incluyendo el primer muestreo completo de Troglodytidae, y utilizamos esta filogenia para inferir sus historias biogeográficas y de diversificación. También inferimos una filogenia expandida que incluye cerca de 100 filoespecies putativas anteriormente documentadas en la literatura, y encontramos que incluir esta diversidad tuvo impactos notables en la historia evolutiva inferida de Certhioidea. Este árbol a nivel de filoespecies documentó algunos casos de parafilia de especies, algunos previamente descritos en la literatura y otros novedosos. Encontramos que Certhioidea se originó principalmente en Eurasia y se dispersó en América del Norte cinco veces en los últimos 20 millones de años, incluyendo en el origen a los “certhioides del Nuevo Mundo,” troglodítidos y atrapamoscas, un clado llamado Orthourae en este estudio. Después de este evento inicial de dispersión, tanto los troglodítidos como los atrapamoscas se diversificaron ampliamente en el hemisferio, con ambos linajes cruzando repetidamente entre continentes. Sin embargo, no detectamos un impacto notable de la formación del Istmo de Panamá en la frecuencia de eventos de dispersión entre América del Norte y del Sur. La inclusión de filoespecies alteró esta inferencia biogeográfica en algunas partes del árbol, pero en general fue en gran medida consistente. Con el muestreo a nivel de especie, encontramos que las tasas de diversificación dentro de Certhioidea fueron en gran medida constantes a lo largo del tiempo, con una desaceleración detectable hacia el presente. En contraste, el muestreo a nivel de filoespecie recuperó una historia de diversificación diferente, con un aumento significativo en la tasa en el nodo de la corona de Orthourae después de dispersarse en las Américas, y tasas de especiación aumentadas especialmente en los géneros Polioptila y Henicorhina. Esta filogenia, en gran medida resuelta, para Certhioidea, ha proporcionado perspectivas importantes sobre la historia evolutiva de este grupo y brinda un marco para futuros trabajos comparativos en este fascinante clado.
KEYWORDS: acoustic signals, behavioral specimens, bird song, data archiving, Open data, reproducibility, taxonomy, vocalizations, archivo de datos, canto de aves, datos abiertos, especímenes de comportamiento, reproducibilidad, señales acústicas, Taxonomía, vocalizaciones
A survey of recent taxonomic studies of birds that included acoustic trait analyses reveals that most studies have not archived the sound recordings that support their conclusions, despite the current availability of online, publicly available collections of bird sounds. In addition, bird sound recordings have often been cited without unique accession numbers that permit unambiguous sample identification and in considerably less detail than other types of samples, such as museum specimens or genetic samples. Both this lack of data openness and the way acoustic samples have been cited undermine the methodological rigor that otherwise characterizes many of these studies, and much invaluable biological data are likely to be lost over time if bird sound recordings are not archived in long-term collections. I suggest that these problems can be easily addressed by embracing the open data movement and adopting some best practices that are widely used in other fields. Just as study skins and DNA sequences are required to be deposited in publicly available collections such as natural history museums and the GenBank, respectively, sound recordings used in taxonomic studies with acoustic trait analyses should be archived in publicly available collections as a condition for publication of associated results. Authors of taxonomic studies involving sounds should archive their sound recordings and provide unique accession numbers for sound recordings examined, and journals and reviewers should ensure that authors have done so. By embracing the open data movement, research studying avian acoustic signals is expected to become more transparent, reproducible, and useful.
LAY SUMMARY
While some studies of acoustic trait variation of birds have archived the recordings that support their conclusions, many have not.
Despite being invaluable behavioral specimens that document part of an individual's phenotype, bird sound recordings have not been treated with the same practices in data archiving as other types of specimens.
Proper archiving of bird sound recordings is worthwhile to promote research on avian acoustic signals that is transparent and reproducible, and to ensure that invaluable biological data are not lost over time.
Authors of taxonomic studies involving sounds should archive their sound recordings in long-term, publicly available collections and provide unique accession numbers for sound recordings.
Journals should require authors of taxonomic studies involving sounds to archive their recordings as a condition for publication and to provide unique accession numbers, and reviewers should confirm that authors have done so.
Researchers studying avian acoustic signals should embrace the open data movement to enhance the transparency, reproducibility, and utility of acoustic trait studies.
Una revisión de los recientes estudios taxonómicos de aves que incluyeron análisis de rasgos acústicos revela que la mayoría de los estudios no han archivado las grabaciones de sonido que respaldan sus conclusiones, a pesar de la disponibilidad actual de colecciones en línea, públicamente accesibles, de sonidos de aves. Además, las grabaciones de sonido de aves a menudo se han citado sin números de acceso únicos que permitan una identificación de muestra inequívoca y con mucho menos detalle que otros tipos de muestras, como especímenes de museo o muestras genéticas. Tanto la falta de apertura de los datos como la forma en que se han citado las muestras acústicas socavan el rigor metodológico que caracteriza de otra manera a muchos de estos estudios, y es probable que se pierdan datos biológicos invaluables con el tiempo si las grabaciones de sonido de aves no se archivan en colecciones a largo plazo. Sugiero que estos problemas pueden abordarse fácilmente adscribiendo al movimiento de datos abiertos y siguiendo algunas de las buenas prácticas ampliamente utilizadas en otros campos. Así como se requiere que las pieles de estudio y las secuencias de ADN se depositen en colecciones públicamente disponibles como museos de historia natural y el GenBank, respectivamente, las grabaciones de sonido utilizadas en estudios taxonómicos con análisis de rasgos acústicos deben archivarse en colecciones públicamente disponibles como condición para la publicación de los resultados asociados. Los autores de estudios taxonómicos que involucran sonidos deben archivar sus grabaciones de sonido y proporcionar números de acceso únicos para las grabaciones de sonido examinadas, y las revistas y revisores deben asegurarse de que los autores lo hayan hecho. Así, se espera que la adscripción al movimiento de datos abiertos posibilite que la investigación sobre las señales acústicas de las aves sea más transparente, reproducible y útil.
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