The long shadow of woody encroachment: An integrated approach to modeling grassland songbird habitat.

Animals must track resources over relatively fine spatial and temporal scales, particularly in disturbance-mediated systems like grasslands. Grassland birds respond to habitat heterogeneity by dispersing among sites within and between years, yet we know little about how they make post-dispersal settlement decisions. Many methods exist to quantify the resource selection of mobile taxa, but the habitat data used in these models are frequently not collected at the same location or time that individuals were present. This spatiotemporal misalignment may lead to incorrect interpretations and adverse conservation outcomes, particularly in dynamic systems. To investigate the extent to which spatially and temporally dynamic vegetation conditions and topography drive grassland bird settlement decisions, we integrated multiple data sources from our study site to predict slope, vegetation height, and multiple metrics of vegetation cover at any point in space and time within the temporal and spatial scope of our study. We paired these predictions with avian mark-resight data for 8 years at the Konza Prairie Biological Station in NE Kansas to evaluate territory selection for Grasshopper Sparrows (Ammodramus savannarum), Dickcissels (Spiza americana), and Eastern Meadowlarks (Sturnella magna). Each species selected different types and amounts of herbaceous vegetation cover, but all three species preferred relatively flat areas with less than 6% shrub cover and less than 1% tree cover. We evaluated several scenarios of woody vegetation removal and found that, with a targeted approach, the simulated removal of just one isolated tree in the uplands created up to 14 ha of grassland bird habitat. This study supports growing evidence that small amounts of woody encroachment can fragment landscapes, augmenting conservation threats to grassland systems. Conversely, these results demonstrate that drastic increases in bird habitat area could be achieved through relatively efficient management interventions. The results and approaches reported pave the way for more efficient conservation efforts in grasslands and other systems through spatiotemporal alignment of habitat with animal behaviors and simulated impacts of management interventions.

Systems biology as a framework to understand the physiological and endocrine bases of behavior and its evolution-From concepts to a case study in birds.

Organismal behavior, with its tremendous complexity and diversity, is generated by numerous physiological systems acting in coordination. Understanding how these systems evolve to support differences in behavior within and among species is a longstanding goal in biology that has captured the imagination of researchers who work on a multitude of taxa, including humans. Of particular importance are the physiological determinants of behavioral evolution, which are sometimes overlooked because we lack a robust conceptual framework to study mechanisms underlying adaptation and diversification of behavior. Here, we discuss a framework for such an analysis that applies a "systems view" to our understanding of behavioral control. This approach involves linking separate models that consider behavior and physiology as their own networks into a singular vertically integrated behavioral control system. In doing so, hormones commonly stand out as the links, or edges, among nodes within this system. To ground our discussion, we focus on studies of manakins (Pipridae), a family of Neotropical birds. These species have numerous physiological and endocrine specializations that support their elaborate reproductive displays. As a result, manakins provide a useful example to help imagine and visualize the way systems concepts can inform our appreciation of behavioral evolution. In particular, manakins help clarify how connectedness among physiological systems-which is maintained through endocrine signaling-potentiate and/or constrain the evolution of complex behavior to yield behavioral differences across taxa. Ultimately, we hope this review will continue to stimulate thought, discussion, and the emergence of research focused on integrated phenotypes in behavioral ecology and endocrinology.

Dancing drives evolution of sexual size dimorphism in manakins.

Body size mediates life history, physiology and inter- and intra-specific interactions. Within species, sexes frequently differ in size, reflecting divergent selective pressures and/or constraints. Both sexual selection and differences in environmentally mediated reproductive constraints can drive sexual size dimorphism, but empirically testing causes of dimorphism is challenging. Manakins (Pipridae), a family of Neotropical birds comprising approximately 50 species, exhibit a broad range of size dimorphism from male- to female-biased and are distributed across gradients of precipitation and elevation. Males perform courtship displays ranging from simple hops to complex aerobatic manoeuvres. We tested associations between sexual size dimorphism and (a) agility and (b) environment, analysing morphological, behavioural and environmental data for 22 manakin species in a phylogenetic framework. Sexual dimorphism in mass was most strongly related to agility, with males being lighter than females in species performing more aerial display behaviours. However, wing and tarsus length dimorphism were more strongly associated with environmental variables, suggesting that different sources of selection act on different aspects of body size. These results highlight the strength of sexual selection in shaping morphology-even atypical patterns of dimorphism-while demonstrating the importance of constraints and ecological consequences of body size evolution.

Phenology of tropical understory trees: patterns and correlates.

Reproductive phenologies of plants are constrained by climate in highly seasonal regions. In contrast, plants growing in wet tropical forests are freed from many abiotic constraints, which in canopy tree communities lead to a rich diversity of phenological patterns within and among individuals, species and communities. However, basic descriptions of tropical phenological patterns and the processes that shape them are rare. Here, we document the individual-, population-, and landscape-level phenological patterns of two dominant families of understory woody plants important to avian frugivores, the Melastomataceae and Rubiaceae, along an elevational transect in Costa Rica. The 226 individual plants belonging to 35 species in this study, varied in the number of reproductive bouts/year, and the timing, duration, and synchrony of reproductive stages. This variation was not related to factors related to their interactions with mutualists and antagonists, nor did it appear to be constrained by phylogeny. Diverse phenological patterns among species led to relatively aseasonal patterns at the community and landscape level. Overall, evidence for biotic processes shaping temporal patterns of fruiting phenology was weak or absent. These findings reveal a number of unexplained patterns, and suggest that factors shaping phenology in relatively aseasonal forests operate in idiosyncratic ways at the species level.

Lekking birds in a tropical forest forego sex for migration.

Facultative, partially migratory animals provide a contemporary window into the evolution of migration, offering rare opportunities to examine the life-history trade-offs associated with migration. For the first time, to our knowledge, we describe the nature of these trade-offs, using a lek-breeding tropical bird, the white-ruffed manakin (Corapipo altera). Previous evidence indicated that weather drives post-breeding migration to lower elevations bringing condition-related benefits. Using elevation-sensitive stable isotope measurements and more than 1200 h of behavioural observations, we show that male manakins which migrate incur costs of diminished social status and matings with females the following breeding season. Because migratory tendency depends on inter-annual variation in weather, physical costs of displays and breeding prospects the following year, migratory decisions are subject to both natural and sexual selection, with the outcome of such decisions linked to changing climatic regimes.

Dancing in the Rain: How Do Abiotic Conditions Influence Sexually Selected Behaviors in the White-Ruffed Manakin?

Tropical animals respond to rainfall in population-specific ways. In extremely wet regions, endotherms experience heavy rains as stressors with consequences for behavior and demography. Ultimately, such stressors can affect the relative strength of abiotic selection, reducing the scope for sexual selection, and other biotic sources of selection. We studied population-level differences in the response to rainfall of White-ruffed manakins (Corapipo altera) on the Caribbean slope of Costa Rica, a species having a lek mating system subject to strong sexual selection. Between 2008 and 2013, we studied reproductive behavior in a population inhabiting an extremely wet site; estimates of apparent survival were low, and the turnover of display courts and dominant males was high. Males also engaged in coordinated display, and sub-adult males practiced in the presence of adults. Over three breeding seasons (2017-2019), we studied a population of the same species at a site only 110 km away, but in a location receiving roughly half as much rain. We tested behavioral predictions of three alternative mechanisms-indirect abiotic effects, direct mortality effects, and direct behavioral effects-linking rainfall to sexual selection in these two populations. Data derived from over 4300 h of observations at 105 display sites revealed high interannual variation in nearly all response variables, including turnover of display sites, retention of alpha status, male display behavior, and time females spent assessing male display. Additionally, we detected spatial differences in drivers of display site turnover. Notably, age distribution of males was skewed toward older individuals at the drier location. Based on these findings, we infer that indirect abiotic effects on forest structure leading to display site transience and direct effects of mortality increasing turnover in the male population likely underlie links between rain and the spatial and temporal differences we documented. Our results are consistent with rain constituting an important source of abiotic selection for tropical endotherms and modulating the scope for sexual selection near the extremes of a species' hygric niche.

Storms drive altitudinal migration in a tropical bird.

Although migration is a widespread and taxonomically diverse behaviour, the ecological factors shaping migratory behaviour are poorly understood. Like other montane taxa, many birds migrate along elevational gradients in the tropics. Forty years ago, Alexander Skutch postulated that severe storms could drive birds to migrate downhill. Here, we articulate a novel mechanism that could link storms to mortality risks via reductions in foraging time and provide, to our knowledge, the first tests of this hypothesis in the White-ruffed Manakin (Corapipo altera), a small partially migratory frugivore breeding on the Atlantic slope of Costa Rica. As predicted, variation in rainfall was associated with plasma corticosterone levels, fat stores, plasma metabolites and haematocrit. By collecting data at high and low elevation sites simultaneously, we also found that high-elevation residents were more adversely affected by storms than low elevation migrants. These results, together with striking temporal capture patterns of altitudinal migrants relative to storms, provide, to our knowledge, the first evidence that weather-related risks incurred by species requiring high food intake rates can explain altitudinal migrations of tropical animals. These findings resolve conflicting evidence for and against food limitation being important in the evolution of this behaviour, and highlight how endogenous and exogenous processes influence life-history trade-offs made by individuals in the wild. Because seasonal storms are a defining characteristic of most tropical ecosystems and rainfall patterns will probably change in ensuing decades, these results have important implications for understanding the ecology, evolution and conservation of tropical animals.

Hygric Niches for Tropical Endotherms.

Biotic selective pressures dominate explanations for the evolutionary ecology of tropical endotherms. Yet, abiotic factors, principally precipitation regimes, shape biogeographical and phenological patterns in tropical regions. Despite its importance, we lack a framework for understanding when, why, and how rain affects endotherms. Here, we review how tropical birds and mammals respond to rain at individual, population, and community levels, and propose a conceptual framework to interpret divergent responses. Diverse direct and indirect mechanisms underlie responses to rainfall, including physiological, top-down, and food-related drivers. Our framework constitutes a roadmap for the empirical studies required to understand the consequences of rainfall variability. Identifying the patterns and mechanisms underpinning responses to temporal variation in precipitation is crucial to anticipate consequences of anthropogenic climate change.

Partial migration in birds: tests of three hypotheses in a tropical lekking frugivore.

1. Partially migratory species provide opportunities to understand which ecological factors cause some animals to migrate when others remain resident year round. Partial migration in birds has been explained by the dominance, arrival-time, and body-size hypotheses. 2. Testing these hypotheses has proven difficult due to the similarities of the predictions they make in temperate-breeding long-distance migrants. In tropical altitudinal migrants, however, these hypotheses make different predictions regarding the sex, age, and condition of migrants and residents. 3. Among white-ruffed manakins in Costa Rica, young birds were not more likely to migrate (as predicted by the dominance hypothesis), nor were females more likely to migrate (as predicted by the arrival-time hypothesis). All condition-related variables interacted with sex, together explaining much of the variation in migratory behaviour. 4. I re-articulate the body-size hypothesis in the context of tropical altitudinal bird migration, focusing explicitly on how limited foraging opportunities and differences in individual condition affect fasting ability during torrential rains. Despite ample food, the smallest birds or those stressed by parasites or moult may risk starvation at breeding elevations due to a reduction in foraging time. These results highlight how intrinsic and extrinsic factors may interact to produce observed patterns of within- and among-species variation in migratory behaviour.

Altitudinal migration: ecological drivers, knowledge gaps, and conservation implications.

Animal migration has been the subject of intensive research for more than a century, but most research has focused on long-distance rather than short-distance migration. Altitudinal migration is a form of short-distance migration in which individuals perform seasonal elevational movements. Despite its geographic and taxonomic ubiquity, there is relatively little information about the intrinsic and extrinsic factors that influence altitudinal migratory behaviour. Without this information, it is difficult to predict how rapid environmental changes will affect population viability of altitudinal migrants. To synthesize current knowledge, we compiled literature on altitudinal migration for all studied taxa, and identified the leading hypotheses explaining this behaviour. Studies of animal altitudinal migration cover many taxonomic lineages, with birds being the most commonly studied group. Altitudinal migration occurs in all continents except for Antarctica, but about a third of the literature focused on altitudinal migration in North America. Most research suggests that food and weather are the primary extrinsic drivers of altitudinal migration. In addition, substantial individual-level variation in migratory propensity exists. Individual characteristics that are associated with sex, dominance rank, and body size explain much of the variation in migratory propensity in partially migratory populations, but individual-level correlates are poorly known for most taxa. More research is needed to quantify the effects of habitat loss, habitat fragmentation, and climate change on altitudinal migrants. Demographic studies of individually marked populations would be particularly valuable for advancing knowledge of the cascading effects of environmental change on migratory propensity, movement patterns, and population viability. We conclude our review with recommendations for study designs and modelling approaches that could be used to narrow existing knowledge gaps, which currently hinder effective conservation of altitudinal migratory species.

Why migrate? A test of the evolutionary precursor hypothesis.

The question of why birds migrate is still poorly understood despite decades of debate. Previous studies have suggested that use of edge habitats and a frugivorous diet are precursors to the evolution of migration in Neotropical birds. However, these studies did not explore other ecological correlates of migration and did not control for phylogeny at the species level. We tested the evolutionary precursor hypothesis by examining the extent to which habitat and diet are associated with migratory behavior, using a species-level comparative analysis of the Tyranni. We used both migratory distance and sedentary versus migratory behavior as response variables. We also examined the influences of foraging group size, membership in mixed-species flocks, elevational range, and body mass on migratory behavior. Raw species analyses corroborated some results from studies that put forth the evolutionary precursor hypothesis, but phylogenetically independent contrast analyses highlighted an important interaction between habitat and diet and their roles as precursors to migration. Foraging group size was consistently associated with migratory behavior in both raw species and independent contrast analyses. Our results lead to a resource variability hypothesis that refines the evolutionary precursor hypothesis and reconciles the results of several studies examining precursors to migration in birds.

Pallid bands in feathers and associated stable isotope signatures reveal effects of severe weather stressors on fledgling sparrows.

In August 2013, we observed a high incidence (44%) of synchronous bands of reduced melanin (a type of fault bar we have termed "pallid bands") across the rectrices of juvenile Grasshopper Sparrows (Ammodrammus savannarum) captured near El Reno, Oklahoma. Earlier that year, on May 31, the site was struck by a severe storm which rained hailstones exceeding 5.5 cm diameter and spawned an historic 4.2 km-wide tornado <8 km to the south of the site. We hypothesized that this stressor had induced the pallid bands. An assessment of Grasshopper Sparrow nesting phenology indicated that a large number of nestlings were likely growing tail feathers when the storm hit. The pallid bands were restricted to the distal half of feathers and their widths significantly increased as a function of distance from the tip (i.e., age at formation). We predicted that if stress had caused these pallid bands, then a spike in circulating δ (15)N originating from tissue catabolism during the stress response would have been incorporated into the developing feather. From 18 juveniles captured at the site in August we measured δ (15)N and δ (13)C stable isotope ratios within four to five 0.25-0.40 mg feather sections taken from the distal end of a tail feather; the pallid band, if present, was contained within only one section. After accounting for individual and across-section variation, we found support for our prediction that feather sections containing or located immediately proximal to pallid bands (i.e., the pallid band region) would show significantly higher δ (15)N than sections outside this region. In contrast, the feathers of juveniles with pallid bands compared to normal appearing juveniles showed significantly lower δ (15)N. A likely explanation is that the latter individuals hatched after the May 31 storm and had consumed a trophically-shifted diet relative to juveniles with pallid bands. Considering this, the juveniles of normal appearance were significantly less abundant within our sample relative to expectations from past cohorts (z = - 2.03; p = 0.042) and, in as much, suggested widespread nest losses during the storm. Severe weather events may represent major stressors to ground-nesting birds, especially for recent fledglings. We call for others to exploit opportunities to study the effects of severe weather when these rare but devastating stressors impact established field research sites.

Altitudinal migration in bats: evidence, patterns, and drivers.

Altitudinal migrations are common in all major vertebrate and some invertebrate lineages. Such migrations have important implications for the basic and applied ecology of animals making these movements. The idea that bats make altitudinal migrations has been suggested for nearly a century. However, studies documenting the existence and causes of altitudinal bat migrations are scarce, and are frequently published in the 'grey' literature. For the first time, we comprehensively review the evidence supporting the existence of altitudinal bat migrations worldwide, describe basic patterns of migration in temperate and tropical regions, and articulate and propose tests of hypotheses potentially explaining these migrations. We compiled a list of 50 studies indicative of altitudinal bat migration in 61 species (five families) from 21 countries (four continents). The temporal and spatial patterns of these migrations grouped biogeographically. Temperate bats generally exhibit sex-biased migrations with females inhabiting lower elevations than males during reproductive periods. Although there is less information on tropical bat migration, few studies report sex-biased migration. We compiled hypotheses proposed in the bat and (more extensive) avian literature to provide a list of hypotheses potentially explaining altitudinal bat migrations. These hypotheses rely upon temporal availability of (and competition for) food resources, spatial distribution of geomorphological features suitable for hibernation, sex-related differences in the use of torpor, mating opportunities, and climatic factors that impose direct physiological challenges to survival or that restrict the ability to forage. A more thorough description of the migration patterns of most species will be required to distinguish effectively among these hypotheses. We identify research avenues that would broaden our understanding of bat migration patterns and provide critical information required for effective conservation.

Evolutionary divergence in brain size between migratory and resident birds.

Despite important recent progress in our understanding of brain evolution, controversy remains regarding the evolutionary forces that have driven its enormous diversification in size. Here, we report that in passerine birds, migratory species tend to have brains that are substantially smaller (relative to body size) than those of resident species, confirming and generalizing previous studies. Phylogenetic reconstructions based on Bayesian Markov chain methods suggest an evolutionary scenario in which some large brained tropical passerines that invaded more seasonal regions evolved migratory behavior and migration itself selected for smaller brain size. Selection for smaller brains in migratory birds may arise from the energetic and developmental costs associated with a highly mobile life cycle, a possibility that is supported by a path analysis. Nevertheless, an important fraction (over 68%) of the correlation between brain mass and migratory distance comes from a direct effect of migration on brain size, perhaps reflecting costs associated with cognitive functions that have become less necessary in migratory species. Overall, our results highlight the importance of retrospective analyses in identifying selective pressures that have shaped brain evolution, and indicate that when it comes to the brain, larger is not always better.

Can variation in risk of nest predation explain altitudinal migration in tropical birds?

Migration is among the best studied of animal behaviors, yet few empirical studies have tested hypotheses explaining the ultimate causes of these cyclical annual movements. Fretwell's (1980) hypothesis predicts that if nest predation explains why many tropical birds migrate uphill to breed, then predation risk must be negatively associated with elevation. Data from 385 artificial nests spanning 2,740 m of elevation on the Atlantic slope of Costa Rica show an overall decline in predation with increasing elevation. However, nest predation risk was highest at intermediate elevations (500-650 m), not at lowest elevations. The proportion of nests depredated by different types of predators differed among elevations. These results imply that over half of the altitudinal migrant bird species in this region migrate to safer breeding areas than their non-breeding areas, suggesting that variation in nest predation risk could be an important benefit of uphill migrations of many species.

Phenology of tropical understory trees: patterns and correlates

Reproductive phenologies of plants are constrained by climate in highly seasonal regions. In contrast, plants growing in wet tropical forests are freed from many abiotic constraints, which in canopy tree communities lead to a rich diversity of phenological patterns within and among individuals, species and communities. However, basic descriptions of tropical phenological patterns and the processes that shape them are rare. Here, we document the individual-, population-, and landscape-level phenological patterns of two dominant families of understory woody plants important to avian frugivores, the Melastomataceae and Rubiaceae, along an elevational transect in Costa Rica. The 226 individual plants belonging to 35 species in this study, varied in the number of reproductive bouts/year, and the timing, duration, and synchrony of reproductive stages. This variation was not related to factors related to their interactions with mutualists and antagonists, nor did it appear to be constrained by phylogeny. Diverse phenological patterns among species led to relatively aseasonal patterns at the community and landscape level. Overall, evidence for biotic processes shaping temporal patterns of fruiting phenology was weak or absent. These findings reveal a number of unexplained patterns, and suggest that factors shaping phenology in relatively aseasonal forests operate in idiosyncratic ways at the species level.

En regiones con marcada estacionalidad, los patrones fenológicos de las plantas están limitados por el clima. Por el contrario, las plantas que crecen en bosques húmedos tropicales, no tienen tantas limitaciones abióticas y es por esto que el dosel presenta una diversidad muy rica en los patrones fenológicos de individuos, especies y comunidades. Sin embargo, es muy escasa la información sobre la descripción básica de los patrones fenológicos tropicales y de los procesos que los afectan. En este documento, presentamos los patrones fenológicos, a nivel de individuo, población y paisaje, a lo largo de un transecto altitudinal en Costa Rica, para dos familias dominantes de plantas leñosas (Melastomataceae y Rubiaceae) que son de gran importancia en la dieta de aves frugívoras. En este estudio, las plantas variaron en el número de brotes reproductivos por año, así como en el tiempo, duración y sincronización de la reproducción. Esta variación no estuvo relacionada con interacciones con mutualistas y antagonistas, ni aparentemente estuvo limitada por la filogenia. Los diversos patrones fenológicos de las especies fueron relativamente no estacionales a nivel de comunidad y paisaje. En conjunto, hubo poca evidencia de que los procesos abióticos hayan afectado los patrones fenológicos de fructificación. Estos resultados revelan una serie de patrones sin explicación aparente, y sugieren que los factores que determinan la fenología en los bosques relativamente no estacionales, funcionan de manera idiosincrática a nivel de especie.