Habitat use of Amazonian birds varies by age and foraging guild along a disturbance gradient.

Patterns of habitat use directly influence a species' fitness, yet for many species an individual's age can influence patterns of habitat use. However, in tropical rainforests, which host the greatest terrestrial species diversity, little is known about how age classes of different species use different adjacent habitats of varying quality. We use long-term mist net data from the Amazon rainforest to assess patterns of habitat use among adult, adolescent (teenage) and young understory birds in forest fragments, primary and secondary forest at the Biological Dynamics of Forest Fragments Project in Brazil. Insectivore adults were most common in primary forest, adolescents were equally likely in primary and secondary forest, and all ages were the least common in forest fragments. In contrast to insectivores, frugivores and omnivores showed no differences among all three habitat types. Our results illustrate potential ideal despotic distributions among breeding populations of some guilds of understory birds where adult insectivores may competitively exclude adolescent individuals from primary forest. Secondary forest recovery appears to hold promise as a breeding habitat for frugivore and omnivore species but only as a pre-breeding habitat for insectivores, but as the forest ages, the demographic structure of bird populations should match that of primary forest.

Equivocal support for the climate variability hypothesis within a Neotropical bird assemblage.

The climate variability hypothesis posits that an organism's exposure to temperature variability determines the breadth of its thermal tolerance and has become an important framework for understanding variation in species' susceptibilities to climate change. For example, ectotherms from more thermally stable environments tend to have narrower thermal tolerances and greater sensitivity to projected climate warming. Among endotherms, however, the relationship between climate variability and thermal physiology is less clear, particularly with regard to microclimate variation-small-scale differences within or between habitats. To address this gap, we explored associations between two sources of temperature variation (habitat type and vertical forest stratum) and (1) thermal physiological traits and (2) temperature sensitivity metrics within a diverse assemblage of Neotropical birds (n = 89 species). We used long-term temperature data to establish that daily temperature regimes in open habitats and forest canopy were both hotter and more variable than those in the forest interior and forest understory, respectively. Despite these differences in temperature regime, however, we found little evidence that species' thermal physiological traits or temperature sensitivity varied in association with either habitat type or vertical stratum. Our findings provide two novel and important insights. First, and in contrast to the supporting empirical evidence from ectotherms, the thermal physiology of birds at our study site appears to be largely decoupled from local temperature variation, providing equivocal support for the climate variability hypothesis in endotherms. Second, we found no evidence that the thermal physiology of understory forest birds differed from that of canopy or open-habitat species-an oft-invoked, yet previously untested, mechanism for why these species are so vulnerable to environmental change.

Harmonizing spatial scales and ecological theories to predict avian richness and functional diversity within forest ecosystems.

Classic ecological theory has proven that temperature, precipitation and productivity organize ecosystems at broad scales and are generalized drivers of biodiversity within different biomes. At local scales, the strength of these predictors is not consistent across different biomes. To better translate these theories to localized scales, it is essential to determine the links between drivers of biodiversity. Here we harmonize existing ecological theories to increase the predictive power for species richness and functional diversity. We test the relative importance of three-dimensional habitat structure as a link between local and broad-scale patterns of avian richness and functional diversity. Our results indicate that habitat structure is more important than precipitation, temperature and elevation gradients for predicting avian species richness and functional diversity across different forest ecosystems in North America. We conclude that forest structure, influenced by climatic drivers, is essential for predicting the response of biodiversity with future shifts in climatic regimes.

Effects of forest fragmentation on avian breeding activity.

Biodiversity declines and ecosystem decay follow forest fragmentation; initially, abundant species may become rare or be extirpated. Underlying mechanisms behind delayed extirpation of certain species following forest fragmentation are unknown. Species declines may be attributed to an inadequate number of breeding adults required to replace the population or decreased juvenile survival rate due to reduced recruitment or increased nest predation pressures. We used 10 years of avian banding data, 5 years before and 4 years after fragment isolation, from the Biological Dynamics of Forest Fragments Project, carried out near Manaus, Brazil, to investigate the breeding activity hypothesis that there is less breeding activity and fewer young after relative to before fragment isolation. We compared the capture rates of active breeding and young birds in 3 forest types (primary forest, fragment before isolation, and fragment after isolation) and the proportion of active breeding and young birds with all birds in each unique fragment type before and after isolation. We grouped all bird species by diet (insectivore or frugivore) and nesting strategy (open cup, cavity, or enclosed) to allow further comparisons among forest types. We found support for the breeding activity hypothesis in insectivorous and frugivorous birds (effect sizes 0.45 and 0.53, respectively) and in birds with open-cup and enclosed nesting strategies (effect sizes 0.56 and 0.44, respectively) such that on average there were more breeding birds in fragments before isolation relative to after isolation. A larger proportion of birds in the community were actively breeding before fragment isolation (72%) than after fragment isolation (11%). Unexpectedly, there was no significant decrease in the number of young birds after fragment isolation, although sample sizes for young were small (n = 43). This may have been due to sustained immigration of young birds to fragments after isolation. Together, our results provide some of the strongest evidence to date that avian breeding activity decreases in response to fragment isolation, which could be a fundamental mechanism contributing to ecosystem decay.

Efectos de la fragmentación del bosque sobre la actividad reproductiva de las aves Resumen Les declinaciones de la biodiversidad y el deterioro de los ecosistemas van después de la fragmentación forestal; al inicio, las especies abundantes pueden volverse raras o ser extirpadas. Todavía no se conocen los mecanismos subyacentes detrás de la extirpación retrasada de ciertas especies después de la fragmentación forestal. La declinación de las especies puede atribuirse a un número inadecuado de adultos reproductivos requeridos para reemplazar a la población o a la tasa reducida de supervivencia de los juveniles debido al reclutamiento disminuido o al incremento en la presión de depredación de los nidos. Usamos diez años de datos de anillamiento de aves, cinco años antes y cuatro años después del aislamiento por fragmentación, tomados del Proyecto Dinámica Biológica de Fragmentos de Bosque realizado cerca de Manaos, Brasil, para investigar la hipótesis de actividad reproductiva que sostiene que hay una menor actividad reproductiva y menos crías después del aislamiento por fragmentación que antes del aislamiento. Comparamos las tasas de captura de aves con reproducción activa y aves juveniles en tres tipos de bosque (primario, fragmento antes del aislamiento y fragmento después del aislamiento) y la proporción de las aves juveniles y con reproducción activa con todas las aves en cada tipo de fragmento único antes y después del aislamiento. Agrupamos todas las especies de aves según su dieta (insectívora o frugívora) y su estrategia de anidación (nido abierto, cavidad o nido cerrado) para poder realizar más comparaciones entre los tipos de bosque. Las aves con dieta insectívora y frugívora (tamaño del efecto: 0.45 y 0.53, respectivamente) y aquellas con nidos abiertos y cerrados (tamaño del efecto: 0.56 y 0.44, respectivamente) respaldaron la hipótesis de la actividad reproductora de tal manera que en promedio hubo más aves reproductoras en los fragmentos antes del aislamiento que después del aislamiento. Una gran parte de las aves de la comunidad tuvieron reproducción activa antes del aislamiento por fragmentación (72%) que después del aislamiento (11%). Sorprendentemente, no hubo una disminución significativa en el número de aves juveniles después del aislamiento, si bien el tamaño de la muestra de este grupo fue reducido (n = 43). Lo anterior pudo deberse a la continua inmigración de juveniles a los fragmentos después del aislamiento. En conjunto, nuestros resultados proporcionan algunos de los indicios más claros de que la actividad reproductiva de las aves disminuye como respuesta al aislamiento por fragmentación, lo cual podría ser un mecanismo fundamental del deterioro de los ecosistemas.

Long-term changes in avian biomass and functional diversity within disturbed and undisturbed Amazonian rainforest.

Recent long-term studies in protected areas have revealed the loss of biodiversity, yet the ramifications for ecosystem health and resilience remain unknown. Here, we investigate how the loss of understory birds, in the lowest stratum of the forest, affects avian biomass and functional diversity in the Amazon rainforest. Across approximately 30 years in the Biological Dynamics of Forest Fragments Project, we used a historical baseline of avian communities to contrast the avian communities in today's primary forest with those in modern disturbed habitat. We found that in primary rainforest, the reduced abundance of insectivorous species led to reduced functional diversity, but no reduction of biomass, indicating that species with similar functional traits are less likely to coexist in modern primary forests. Because today's forests contain fewer functionally redundant species-those with similar traits-we argue that avian communities in modern primary Amazonian rainforests are less resilient, which may ultimately disrupt the ecosystem in dynamic and unforeseen ways.

Maintenance of local adaptation despite gene flow in a coastal songbird.

Adaptation to local environments is common in widespread species and the basis of ecological speciation. The song sparrow (Melospiza melodia) is a widespread, polytypic passerine that occurs in shrubland habitats throughout North America. We examined the population structure of two parapatric subspecies that inhabit different environments: the Atlantic song sparrow (M. m. atlantica), a coastal specialist, and the eastern song sparrow (M. m. melodia), a shrubland generalist. These populations lacked clear mitochondrial population structure, yet coastal birds formed a distinct nuclear genetic cluster. We found weak overall genomic differentiation between these subspecies, suggesting either recent divergence, extensive gene flow, or a combination thereof. There was a steep genetic cline at the transition to coastal habitats, consistent with isolation by environment, not isolation by distance. A phenotype under divergent selection, bill size, varied with the amount of coastal ancestry in transitional areas, but larger bill size was maintained in coastal habitats regardless of ancestry, further supporting a role for selection in the maintenance of these subspecies. Demographic modeling suggested a divergence history of limited gene flow followed by secondary contact, which has emerged as a common theme in adaptive divergence across taxa.

The uncertain future of protected lands and waters.

Protected areas are intended to safeguard biodiversity in perpetuity, yet evidence suggests that widespread legal changes undermine protected area durability and efficacy. We documented these legal changes-protected area downgrading, downsizing, and degazettement (PADDD) events-in the United States and Amazonian countries and compiled available data globally. Governments of the United States and Amazonian countries enacted 269 and 440 PADDD events, respectively. Between 1892 and 2018, 73 countries enacted 3749 PADDD events, removing 519,857 square kilometers from protection and tempering regulations in an additional 1,659,972 square kilometers; 78% of events were enacted since 2000. Most PADDD events (62%) are associated with industrial-scale resource extraction and development, suggesting that PADDD may compromise biodiversity conservation objectives. Strategic policy responses are needed to address PADDD and sustain effective protected areas.

Noise-induced vocal plasticity in urban white-crowned sparrows does not involve adjustment of trill performance components.

Background noise can interfere with acoustic communication and subsequently influence signaling behavior. Immediate signaling flexibility (ISF) is a context-dependent form of behavioral plasticity that allows animals to temporarily change their acoustic behavior in response to noise fluctuations and potentially improve the chances of successful communication in noisy environments. The adaptive value of ISF is ultimately contingent on the response of the intended receiver, and there are differential effects on receiver response depending on which signal component is modified. However, there is scant research on whether ISF involves modification of a signal component specifically linked to mate attraction or territory defense. Our study addresses this knowledge gap and provides important insight into whether males employ short-term signal modification in a manner that could affect mate pairing success in birds. Specifically, we explore the maladaptive potential of ISF in the San Francisco, California population of Zonotrichia leucophrys nuttalli by testing for changes in trill bandwidth and rate-the specific trill structure components known to influence the receiver's perception of vocal performance in this species-before and during noise broadcast experiments. Although Zonotrichia leucophrys nuttalli are capable of ISF, we found no evidence that noise induces temporary adjustment of the trill structure traits used by receivers to assess vocal performance.

White-crowned sparrow males show immediate flexibility in song amplitude but not in song minimum frequency in response to changes in noise levels in the field.

The soundscape acts as a selective agent on organisms that use acoustic signals to communicate. A number of studies document variation in structure, amplitude, or timing of signal production in correspondence with environmental noise levels thus supporting the hypothesis that organisms are changing their signaling behaviors to avoid masking. The time scale at which organisms respond is of particular interest. Signal structure may evolve across generations through processes such as cultural or genetic transmission. Individuals may also change their behavior during development (ontogenetic change) or in real time (i.e., immediate flexibility). These are not mutually exclusive mechanisms, and all must be investigated to understand how organisms respond to selection pressures from the soundscape. Previous work on white-crowned sparrows (Zonotrichia leucophrys) found that males holding territories in louder areas tend to sing higher frequency songs and that both noise levels and song frequency have increased over time (30 years) in urban areas. These previous findings suggest that songs are changing across generations; however, it is not known if this species also exhibits immediate flexibility. Here, we conducted an exploratory, observational study to ask whether males change the minimum frequency of their song in response to immediate changes in noise levels. We also ask whether males sing louder, as increased minimum frequency may be physiologically linked to producing sound at higher amplitudes, in response to immediate changes in environmental noise. We found that territorial males adjust song amplitude but not minimum frequency in response to changes in environmental noise levels. Our results suggest that males do not show immediate flexibility in song minimum frequency, although experimental manipulations are needed to test this hypothesis further. Our work highlights the need to investigate multiple mechanisms of adaptive response to soundscapes.

Determinants of bird conservation-action implementation and associated population trends of threatened species.

Conservation actions, such as habitat protection, attempt to halt the loss of threatened species and help their populations recover. The efficiency and the effectiveness of actions have been examined individually. However, conservation actions generally occur simultaneously, so the full suite of implemented conservation actions should be assessed. We used the conservation actions underway for all threatened and near-threatened birds of the world (International Union for Conservation of Nature Red List of Threatened Species) to assess which biological (related to taxonomy and ecology) and anthropogenic (related to geoeconomics) factors were associated with the implementation of different classes of conservation actions. We also assessed which conservation actions were associated with population increases in the species targeted. Extinction-risk category was the strongest single predictor of the type of conservation actions implemented, followed by landmass type (continent, oceanic island, etc.) and generation length. Species targeted by invasive nonnative species control or eradication programs, ex situ conservation, international legislation, reintroduction, or education, and awareness-raising activities were more likely to have increasing populations. These results illustrate the importance of developing a predictive science of conservation actions and the relative benefits of each class of implemented conservation action for threatened and near-threatened birds worldwide.

Patterns of Song across Natural and Anthropogenic Soundscapes Suggest That White-Crowned Sparrows Minimize Acoustic Masking and Maximize Signal Content.

Soundscapes pose both evolutionarily recent and long-standing sources of selection on acoustic communication. We currently know more about the impact of evolutionarily recent human-generated noise on communication than we do about how natural sounds such as pounding surf have shaped communication signals over evolutionary time. Based on signal detection theory, we hypothesized that acoustic phenotypes will vary with both anthropogenic and natural background noise levels and that similar mechanisms of cultural evolution and/or behavioral flexibility may underlie this variation. We studied song characteristics of white-crowned sparrows (Zonotrichia leucophrys nuttalli) across a noise gradient that includes both anthropogenic and natural sources of noise in San Francisco and Marin counties, California, USA. Both anthropogenic and natural soundscapes contain high amplitude low frequency noise (traffic or surf, respectively), so we predicted that birds would produce songs with higher minimum frequencies in areas with higher amplitude background noise to avoid auditory masking. We also anticipated that song minimum frequencies would be higher than the projected lower frequency limit of hearing based on site-specific masking profiles. Background noise was a strong predictor of song minimum frequency, both within a local noise gradient of three urban sites with the same song dialect and cultural evolutionary history, and across the regional noise gradient, which encompasses 11 urban and rural sites, several dialects, and several anthropogenic and natural sources of noise. Among rural sites alone, background noise tended to predict song minimum frequency, indicating that urban sites were not solely responsible for driving the regional pattern. These findings support the hypothesis that songs vary with local and regional soundscapes regardless of the source of noise. Song minimum frequency from five core study sites was also higher than the lower frequency limit of hearing at each site, further supporting the hypothesis that songs vary to transmit through noise in local soundscapes. Minimum frequencies leveled off at noisier sites, suggesting that minimum frequencies are constrained to an upper limit, possibly to retain the information content of wider bandwidths. We found evidence that site noise was a better predictor of song minimum frequency than territory noise in both anthropogenic and natural soundscapes, suggesting that cultural evolution rather than immediate behavioral flexibility is responsible for local song variation. Taken together, these results indicate that soundscapes shape song phenotype across both evolutionarily recent and long-standing soundscapes.

Production and perception of communicatory signals in a noisy environment.

Many animals communicate in situations that make it difficult to discriminate a species' signals from those of others. Consequently, coexisting species usually have signals that differ by more than the minimum required to prevent overlap in acoustic features. These gaps between signals might facilitate detection and discrimination of degraded signals in noisy natural conditions. If so, perception of signals should have broader scope than production. We investigated this possibility by studying song production and perception of two species of birds in an especially noisy environment, the Amazonian dawn chorus. With software developed for this study, we digitally synthesized songs of two species, as well as intermediate versions of their songs. Experimental playbacks of these synthesized songs to individuals of both species confirmed that perception (as indicated by responses) was broader than production of songs. We propose that broader perception than production of song promotes communication in noisy situations and limits the similarity between signals of coexisting species.

Signaller: receiver coordination and the timing of communication in Amazonian birds.

The efficacy of communication relies on the detection of signals against background noise. Some species are known to alter the timing of vocalizations to avoid acoustic interference from similar signals of other species, but nothing is known about the possibility of coordinated adjustments in the timing of receivers' attention. I examined the possibility that co-occurring species might respond as well as vocalize at different times in a diverse tropical avifauna by presenting playbacks of recordings to territorial birds at typical and atypical times for singing during the dawn chorus. The results show that co-occurring species of birds in a diverse avifauna partition the timing of both production and response in a way that would reduce acoustic interference between species.