Precipitation explains Wood Thrush (Hylocichla mustelina) occupancy patterns in Northern Costa Rica / Patrones de ocurrencia del Zorzal del Bosque (Hylocichla mustelina) explicados por la precipitación en el Norte de Costa Rica

Abstract Introduction: The Wood Thrush is a migratory bird that has experienced dramatic declines in its populations in recent decades. This species overwinters in forest fragments with intermediate levels of habitat modification in Central America. However, more studies detailing the use of remnant forests through time are needed to elucidate the threats this species faces in the wintering grounds. Objective: To understand the effects of environmental and forest structure variables on the occupancy of Wood Thrush in Northern Costa Rica. Methods: The study area was the Área de Conservación Guanacaste (ACG), located in Northern Costa Rica, in December 2016, and during the 2018-2019 migration season. We estimated Wood Thrush occupancy and detection probability in four locations of ACG (dry forest, cloud forest, and two locations in the wet forest) using single-season occupancy models. We also estimated Wood Thrush occupancy and probability of persistence in different months in three vegetation types (open area, secondary forest, and old-growth forest) in the wet forest of ACG using a multi-season occupancy model approach. Results: Wood Thrush occupancy was best described by precipitation in the four locations of the ACG; the probability of occupancy increased with precipitation. The average occupancy of Wood Thrushes varied with vegetation type: open area with shrubs and forest edge (0.69 ± 0.09), secondary forest (0.46 ± 0.1), and old-growth forest (0.61 ± 0.1). Wood Thrush probability of persistence responded partially to changes in precipitation, with an unexpected increase in persistence when the rainfall continued decreasing in the season. Conclusion: Wood Thrush occupancy was best predicted by changes in precipitation considering a larger spatial scale. Its probability of persistence partially varied with precipitation. An increase in persistence closer to Spring migration might be explained by the start of the breeding season of resident birds, potentially reducing territorial conflicts and conserving energy before migration. The long-term protection of wet forests in Northern Costa Rica is of paramount importance for the conservation of Wood Thrushes in their wintering grounds.

Resumen Introducción: El Zorzal del Bosque es un ave migratoria que ha experimentado caídas dramáticas en sus poblaciones en las últimas décadas. Esta especie pasa el invierno en fragmentos de bosque con niveles intermedios de modificación de hábitat en Centroamérica. Sin embargo, se necesitan más estudios que detallen el uso de los bosques remanentes a lo largo del tiempo para dilucidar las amenazas que enfrenta esta especie en las zonas de invernada. Objetivo: Comprender los efectos de variables ambientales y de estructura del bosque en la ocurrencia del Zorzal del Bosque en el Norte de Costa Rica. Métodos: El área de estudio fue el Área de Conservación Guanacaste (ACG), ubicada en el Norte de Costa Rica, en diciembre de 2016, y en la temporada migratoria 2018-2019. Estimamos la ocurrencia y la probabilidad de detección del Zorzal del Bosque en cuatro ubicaciones de ACG (bosque seco, bosque nuboso y dos ubicaciones en el bosque húmedo) utilizando modelos de ocurrencia de una sola temporada. También estimamos la ocurrencia del Zorzal del Bosque y la probabilidad de persistencia en diferentes meses en tres tipos de vegetación (área abierta, bosque secundario y bosque primario) en el bosque húmedo de ACG utilizando un enfoque de modelo de ocurrencia multi-estacional. Resultados: La ocurrencia del Zorzal del Bosque estuvo mejor descrita por la precipitación en las cuatro localidades del ACG; la probabilidad de ocurrencia aumentó con las precipitaciones. La ocurrencia media de zorzales varió con el tipo de vegetación: área abierta con arbustos y borde de bosque (0.69 ± 0.09), bosque secundario (0.46 ± 0.1) y bosque primario (0.61 ± 0.1). La probabilidad de persistencia del zorzal respondió parcialmente a cambios en la precipitación, con un aumento inesperado en la persistencia cuando las precipitaciones continuaron disminuyendo en la temporada. Conclusión: La ocurrecia del Zorzal del Bosque varió con la precipitación considerando una escala espacial mayor. Su probabilidad de persistencia varió parcialmente con la precipitación. Un aumento en la persistencia más cerca de la migración de primavera podría explicarse por el inicio de la temporada de reproducción de las aves residentes, lo que podría reducir los conflictos territoriales y conservar energía antes de la migración. La protección a largo plazo de los bosques húmedos en el norte de Costa Rica es de suma importancia para la conservación de los Zorzales del Bosque en sus zonas de invernada.

Song plasticity over time and vocal learning in clay-colored thrushes.

Songbirds have been traditionally classified into close-ended or open-ended learning species according to the length of the sensitive period during which birds are able to memorize new vocalizations. Closed-ended learners are generally not capable of changing their song after the first year of life, while open-ended learners show song plasticity as adults. A few Turdus species have been suggested to be open-ended learners, but no long-term study has been conducted to investigate their song plasticity over time. We analyzed the songs of clay-colored thrushes, T. grayi, over four successive breeding seasons to assess song plasticity in their syllable repertoires within and between breeding seasons. A total of 16,262 syllables were classified through visual inspection of spectrograms and multidimensional scaling analysis based on spectrogram correlations. On average, 563 ± 153 (SD) syllables per male per breeding season were analyzed. Male repertoire size was 9-20 syllable types. Males were capable of modifying their syllable repertoire between the initial and final periods of the breeding season. Song plasticity within breeding seasons may be associated with imitation between neighboring males, suggesting song learning in males that were ≥2 years old. This short-term plasticity is not enough, however, to explain the high proportion of change (mean = 65 % syllable types) in repertoire composition between breeding seasons in adult males. Song plasticity resulting from annual changes in repertoire composition could be explained by open-ended learning, but another mechanism, extended memory and re-expression, could also explain long-term plasticity. Experimental studies controlling the acoustic environment are needed to determine which mechanism is responsible for such a high level of song plasticity.