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.

Integrating data types to estimate spatial patterns of avian migration across the Western Hemisphere.

For many avian species, spatial migration patterns remain largely undescribed, especially across hemispheric extents. Recent advancements in tracking technologies and high-resolution species distribution models (i.e., eBird Status and Trends products) provide new insights into migratory bird movements and offer a promising opportunity for integrating independent data sources to describe avian migration. Here, we present a three-stage modeling framework for estimating spatial patterns of avian migration. First, we integrate tracking and band re-encounter data to quantify migratory connectivity, defined as the relative proportions of individuals migrating between breeding and nonbreeding regions. Next, we use estimated connectivity proportions along with eBird occurrence probabilities to produce probabilistic least-cost path (LCP) indices. In a final step, we use generalized additive mixed models (GAMMs) both to evaluate the ability of LCP indices to accurately predict (i.e., as a covariate) observed locations derived from tracking and band re-encounter data sets versus pseudo-absence locations during migratory periods and to create a fully integrated (i.e., eBird occurrence, LCP, and tracking/band re-encounter data) spatial prediction index for mapping species-specific seasonal migrations. To illustrate this approach, we apply this framework to describe seasonal migrations of 12 bird species across the Western Hemisphere during pre- and postbreeding migratory periods (i.e., spring and fall, respectively). We found that including LCP indices with eBird occurrence in GAMMs generally improved the ability to accurately predict observed migratory locations compared to models with eBird occurrence alone. Using three performance metrics, the eBird + LCP model demonstrated equivalent or superior fit relative to the eBird-only model for 22 of 24 species-season GAMMs. In particular, the integrated index filled in spatial gaps for species with over-water movements and those that migrated over land where there were few eBird sightings and, thus, low predictive ability of eBird occurrence probabilities (e.g., Amazonian rainforest in South America). This methodology of combining individual-based seasonal movement data with temporally dynamic species distribution models provides a comprehensive approach to integrating multiple data types to describe broad-scale spatial patterns of animal movement. Further development and customization of this approach will continue to advance knowledge about the full annual cycle and conservation of migratory birds.

Morphological consequences of climate change for resident birds in intact Amazonian rainforest.

Warming from climate change is expected to reduce body size of endotherms, but studies from temperate systems have produced equivocal results. Over four decades, we collected morphometric data on a nonmigratory understory bird community within Amazonian primary rainforest that is experiencing increasingly extreme climate. All 77 species showed lower mean mass since the early 1980s­nearly half with 95% confidence. A third of species concomitantly increased wing length, driving a decrease in mass:wing ratio for 69% of species. Seasonal precipitation patterns were generally better than temperature at explaining morphological variation. Short-term climatic conditions affected all metrics, but time trends in wing and mass:wing remained robust even after controlling for annual seasonal conditions. We attribute these results to pressures to increase resource economy under warming. Both seasonal and long-term morphological shifts suggest response to climate change and highlight its pervasive consequences, even in the heart of the world's largest rainforest.

The interplay of wind and uplift facilitates over-water flight in facultative soaring birds.

Flying over the open sea is energetically costly for terrestrial birds. Despite this, over-water journeys of many birds, sometimes hundreds of kilometres long, are uncovered by bio-logging technology. To understand how these birds afford their flights over the open sea, we investigated the role of atmospheric conditions, specifically wind and uplift, in subsidizing over-water flight at a global scale. We first established that ΔT, the temperature difference between sea surface and air, is a meaningful proxy for uplift over water. Using this proxy, we showed that the spatio-temporal patterns of sea-crossing in terrestrial migratory birds are associated with favourable uplift conditions. We then analysed route selection over the open sea for five facultative soaring species, representative of all major migratory flyways. The birds maximized wind support when selecting their sea-crossing routes and selected greater uplift when suitable wind support was available. They also preferred routes with low long-term uncertainty in wind conditions. Our findings suggest that, in addition to wind, uplift may play a key role in the energy seascape for bird migration that in turn determines strategies and associated costs for birds crossing ecological barriers such as the open sea.

Long-term change in the avifauna of undisturbed Amazonian rainforest: ground-foraging birds disappear and the baseline shifts.

How are rainforest birds faring in the Anthropocene? We use bird captures spanning > 35 years from 55 sites within a vast area of intact Amazonian rainforest to reveal reduced abundance of terrestrial and near-ground insectivores in the absence of deforestation, edge effects or other direct anthropogenic landscape change. Because undisturbed forest includes far fewer terrestrial and near-ground insectivores than it did historically, today's fragments and second growth are more impoverished than shown by comparisons with modern 'control' sites. Any goals for bird community recovery in Amazonian second growth should recognise that a modern bird community will inevitably differ from a baseline from > 35 years ago. Abundance patterns driven by landscape change may be the most conspicuous manifestation of human activity, but biodiversity declines in undisturbed forest represent hidden losses, possibly driven by climate change, that may be pervasive in intact Amazonian forests and other systems considered to be undisturbed.

Juvenile Osprey Navigation during Trans-Oceanic Migration.

To compensate for drift, an animal migrating through air or sea must be able to navigate. Although some species of bird, fish, insect, mammal, and reptile are capable of drift compensation, our understanding of the spatial reference frame, and associated coordinate space, in which these navigational behaviors occur remains limited. Using high resolution satellite-monitored GPS track data, we show that juvenile ospreys (Pandion haliaetus) are capable of non-stop constant course movements over open ocean spanning distances in excess of 1500 km despite the perturbing effects of winds and the lack of obvious landmarks. These results are best explained by extreme navigational precision in an exogenous spatio-temporal reference frame, such as positional orientation relative to Earth's magnetic field and pacing relative to an exogenous mechanism of keeping time. Given the age (<1 year-old) of these birds and knowledge of their hatching site locations, we were able to transform Enhanced Magnetic Model coordinate locations such that the origin of the magnetic coordinate space corresponded with each bird's nest. Our analyses show that trans-oceanic juvenile osprey movements are consistent with bicoordinate positional orientation in transformed magnetic coordinate or geographic space. Through integration of movement and meteorological data, we propose a new theoretical framework, chord and clock navigation, capable of explaining the precise spatial orientation and temporal pacing performed by juvenile ospreys during their long-distance migrations over open ocean.

Understory bird communities in Amazonian rainforest fragments: species turnover through 25 years post-isolation in recovering landscapes.

Inferences about species loss following habitat conversion are typically drawn from short-term surveys, which cannot reconstruct long-term temporal dynamics of extinction and colonization. A long-term view can be critical, however, to determine the stability of communities within fragments. Likewise, landscape dynamics must be considered, as second growth structure and overall forest cover contribute to processes in fragments. Here we examine bird communities in 11 Amazonian rainforest fragments of 1-100 ha, beginning before the fragments were isolated in the 1980s, and continuing through 2007. Using a method that accounts for imperfect detection, we estimated extinction and colonization based on standardized mist-net surveys within discreet time intervals (1-2 preisolation samples and 4-5 post-isolation samples). Between preisolation and 2007, all fragments lost species in an area-dependent fashion, with loss of as few as <10% of preisolation species from 100-ha fragments, but up to 70% in 1-ha fragments. Analysis of individual time intervals revealed that the 2007 result was not due to gradual species loss beginning at isolation; both extinction and colonization occurred in every time interval. In the last two samples, 2000 and 2007, extinction and colonization were approximately balanced. Further, 97 of 101 species netted before isolation were detected in at least one fragment in 2007. Although a small subset of species is extremely vulnerable to fragmentation, and predictably goes extinct in fragments, developing second growth in the matrix around fragments encourages recolonization in our landscapes. Species richness in these fragments now reflects local turnover, not long-term attrition of species. We expect that similar processes could be operating in other fragmented systems that show unexpectedly low extinction.

Dispersal of Amazonian birds in continuous and fragmented forest.

Many ecologists believe birds disappear from tropical forest fragments because they are poor dispersers. We test this idea using a spatially explicit capture data base from the Biological Dynamics of Forest Fragments Project near Manaus, Brazil. We measure bird movements directly, over relatively large scales of space and time, both before and after landscape fragmentation. We found that species which disappear from fragments move extensively between plots before isolation, but not after, and often disperse to longer distances in continuous forest than in fragmented forest. Such species also preferentially emigrate from smaller to larger fragments, showing no preference in continuous forest. In contrast, species that persist in fragments are generally less mobile, do not cross gaps as often, yet disperse further after fragmentation than before. 'Heavy tailed' probability models usually explain dispersal kernels better than exponential or Gaussian models, suggesting tropical forest birds may be better dispersers than assumed with some individuals moving very long distances.

A large-scale deforestation experiment: effects of patch area and isolation on Amazon birds.

As compared with extensive contiguous areas, small isolated habitat patches lack many species. Some species disappear after isolation; others are rarely found in any small patch, regardless of isolation. We used a 13-year data set of bird captures from a large landscape-manipulation experiment in a Brazilian Amazon forest to model the extinction-colonization dynamics of 55 species and tested basic predictions of island biogeography and metapopulation theory. From our models, we derived two metrics of species vulnerability to changes in isolation and patch area. We found a strong effect of area and a variable effect of isolation on the predicted patch occupancy by birds.

Long-term landscape change and bird abundance in Amazonian rainforest fragments.

The rainforests of the Amazon basin are being cut by humans at a rate >20,000 km2/year leading to smaller and more isolated patches of forest, with remaining fragments often in the range of 1-100 ha. We analyzed samples of understory birds collected over 20 years from a standardized mist-netting program in 1- to 100-ha rainforest fragments in a dynamic Amazonian landscape near Manaus, Brazil. Across bird guilds, the condition of second growth immediately surrounding fragments was often as important as fragment size or local forest cover in explaining variation in abundance. Some fragments surrounded by 100 m of open pasture showed reductions in insectivorous bird abundance of over 95%, even in landscapes dominated by continuous forest and old second growth. These extreme reductions may be typical throughout Amazonia in small (< or =10 ha), isolated fragments of rainforest. Abundance for some guilds returned to preisolation levels in 10- and 100-ha fragments connected to continuous forest by 20-year-old second growth. Our results show that the consequences of Amazonian forest loss cannot be accurately described without explicit consideration of vegetation dynamics in matrix habitat. Any dichotomous classification of the landscape into 'forest" and "nonforest" misses essential information about the matrix.

Comparison of wing morphology in three birds of prey: correlations with differences in flight behavior.

Flight is the overriding characteristic of birds that has influenced most of their morphological, physiological, and behavioral features. Flight adaptations are essential for survival in the wide variety of environments that birds occupy. Therefore, locomotor structure, including skeletal and muscular characteristics, is adapted to reflect the flight style necessitated by different ecological niches. Red-tailed hawks (Buteo jamaicensis) soar to locate their prey, Cooper's hawks (Accipiter cooperii) actively chase down avian prey, and ospreys (Pandion haliaetus) soar and hover to locate fish. In this study, wing ratios, proportions of skeletal elements, and relative sizes of selected flight muscles were compared among these species. Oxidative and glycolytic enzyme activities of several muscles were also analyzed via assays for citrate synthase (CS) and for lactate dehydrogenase (LDH). It was found that structural characteristics of these three raptors differ in ways consistent with prevailing aerodynamic models. The similarity of enzymatic activities among different muscles of the three species shows low physiological differentiation and suggests that wing architecture may play a greater role in determining flight styles for these birds.

Rates of species loss from Amazonian forest fragments.

In the face of worldwide habitat fragmentation, managers need to devise a time frame for action. We ask how fast do understory bird species disappear from experimentally isolated plots in the Biological Dynamics of Forest Fragments Project, central Amazon, Brazil. Our data consist of mist-net records obtained over a period of 13 years in 11 sites of 1, 10, and 100 hectares. The numbers of captures per species per unit time, analyzed under different simplifying assumptions, reveal a set of species-loss curves. From those declining numbers, we derive a scaling rule for the time it takes to lose half the species in a fragment as a function of its area. A 10-fold decrease in the rate of species loss requires a 1,000-fold increase in area. Fragments of 100 hectares lose one half of their species in <15 years, too short a time for implementing conservation measures.

Effects of Forest Fragmentation on Understory Hummingbirds in Amazonian Brazil.

We sampled understory bummingbirds in Amazonian forest fragments from before isolation through nine years after isolation. We recorded 377 captures of eight species in five 1-ba fragments and four 10-ha fragments. The three species netted before isolation, Phaethornis superciliosus, Phaethornis bourcieri, and Thalurania furcata, were nearly equally abundant at that time. After isolation abundance of P. bourcieri and T. furcata did not change, but P. superciliosus became nearly twice as common. Five additional species that were netted only after isolation represented about 10% of the post-isolation sample. The species recorded only after isolation were forest species usually found above the levels of nets; fragments were not colonized by nonforest species. Use of fragments did not differ between 1-and 10-ba fragments. The landscape surrounding the fragments included active cattle pasture, abandoned pasture, and Cecropia-dominated second growth, but this variation bad little effect on use of fragments by hummingbirds. The results suggest that these understory hummingbirds can persist in a matrix of fragments, secondary growth, and large forest patches. This response is much different than that of the insectivorous birds that dominate the understory bird community at the site, which are much more vulnerable to fragmentation. Efectos de la fragmentación del bosque sobre los colibrís de sotobosque en la Amazonia Brasileña.

Resumen: Nosotros hemos tomado muestras de colibrís de sotobosque en fragmentos forestales Amazónicos desde antes del aislamiento a través de nueve años después del aislamiento. Registramos 377 capturas de ocho especies en cinco fragmentos de 1-ha, y cuatro fragmentos de 10-ha. Las tres especies capturadas antes del aislamiento, Phaethornis superciliosus, Phaethornis bourcieri, y Thalurania furcata, presentaron casi la misma abundancia en esa época. Después del aislamiento, la abundancia de Phaethornis bourcieri y Thalurania furcata no cambió, pero Phaethornis superciliosus se hizo dos veces más común. Cinco especies más que fueron capturadas solamente después del aislamiento representaron aproximadamente el 10% de la muestra de post aislamiento. Las especies capturadas solamente después del aislamiento eran especies del bosque, las que generalmente se encuentran encima del nivel de las redes; los fragmentos no fueron colonizados por especies no-florestales. La utilización de fragmentos no se diferenció entre los fragmentos de 1- y 10-ha. El paisaje alrededor de los fragmentos incluía pasto activo de ganado vacuno, pasto abandonado, y crecimiento secundario dominado por Cecropia, pero esta variación no afectó mucho la utilización de fragmentos por colibrís. Estos resultados sugieren que estas especies de colibrís de sotobosque pueden persistir en una matriz de fragmentos, crecimiento secundario, y parches de bosque de gran tamaño. Esta reacción es muy diferente a la de los pájaros insectívoros que dominan la comunidad de pájaros del sotohosque en nuestro sitio, los cuales son mucho más vulnerables a la fragmentación.