Publications

Climate change is expected to increase the prevalence and severity of extreme weather events like drought. For many opportunistically breeding tropical bird species, precipitation serves as a primary cue for onset of breeding, and sustained drought can have major impacts on reproductive fitness. The physiological effects of drought are poorly understood, but understanding physiological responses can help resolve the mechanisms underlying effects of drought on population demography. We used physiological data collected on a year-round breeding New Guinea endemic passerine, the White-shouldered Fairywren (Malurus alboscapulatus), during years with persistent rainfall and during the 2015–2016 El Niño drought event to assess patterns of breeding readiness, body condition, and molt. Many males lost their cloacal protuberances, a sperm storage organ they typically maintain year-round, during the drought period. Body molt, which also occurs year-round in this species, was higher in males during the drought period. Using a sliding-window statistical model approach, we identified a critical precipitation window of 0–40 days prior to capture for cloacal protuberance volume and a window of 8–53 days for molt, with lower precipitation associated with smaller cloacal protuberances and greater molt. Plasma androgens were predictive of variation in male cloacal protuberance volume and molt, thus potentially mediating transitions between life-history stages in response to environmental conditions. Female molt, androgens, and male mass varied with photoperiod, with longer but decreasing day lengths characteristic of the austral summer associated with increasing molt and decreasing androgens. Collectively, our results indicate the potential for photoperiod to cue some life-history stages, whereas reduced rainfall impacts male but apparently not female reproductive physiology and readiness to breed. Improving our understanding of environmental regulation of life-history transitions in tropical taxa is essential for identifying which taxa are most at risk under a changing climate.

Seasonally breeding animals often exhibit different social structures during non-breeding and breeding periods that coincide with seasonal environmental variation and resource abundance. However, we know little about the environmental factors associated with when seasonal shifts in social structure occur. This lack of knowledge contrasts with our well-defined knowledge of the environmental cues that trigger a shift to breeding physiology in seasonally breeding species. Here, we identified some of the main environmental factors associated with seasonal shifts in social structure and initiation of breeding in the red-backed fairywren (Malurus melanocephalus), an Australian songbird. Social network analyses revealed that social groups, which are highly territorial during the breeding season, interact in social “communities” on larger home ranges during the non-breeding season. Encounter rates among non-breeding groups were related to photoperiod and rainfall, with shifting photoperiod and increased rainfall associated with a shift toward territorial breeding social structure characterized by reductions in home range size and fewer encounters among nonbreeding social groups. Similarly, onset of breeding was highly seasonal and was also associated with non-breeding season rainfall, with greater rainfall leading to earlier breeding. These findings reveal that for some species, the environmental factors associated with the timing of shifts in social structure across seasonal boundaries can be similar to those that determine timing of breeding. This study increases our understanding of the environmental factors associated with seasonal variation in social structure and how the timing of these shifts may respond to changing climates.

Seed and pollen dispersal contribute to gene flow and shape the genetic patterns of plants over fine spatial scales. We inferred fine-scale spatial genetic structure (FSGS) and estimated realized dispersal distances in Phytelephas aequatorialis, a Neotropical dioecious large-seeded palm. We aimed to explore how seed and pollen dispersal shape this genetic pattern in a focal population. For this purpose, we genotyped 138 seedlings and 99 adults with 20 newly developed microsatellite markers. We tested if rodent-mediated seed dispersal has a stronger influence than insect-mediated pollen dispersal in shaping FSGS. We also tested if pollen dispersal was influenced by the density of male palms around mother palms in order to further explore this ecological process in large-seeded plants. Rodent-mediated dispersal of these large seeds occurred mostly over short distances (mean 34.76 ± 34.06 m) while pollen dispersal distances were two times higher (mean 67.91 ± 38.29 m). The spatial extent of FSGS up to 35 m and the fact that seed dispersal did not increase the distance at which male alleles disperse suggest that spatially limited seed dispersal is the main factor shaping FSGS and contributes only marginally to gene flow within the population. Pollen dispersal distances depended on the density of male palms, decreasing when individuals show a clumped distribution and increasing when they are scattered. Our results show that limited seed dispersal mediated by rodents shapes FSGS in P. aequatorialis, while more extensive pollen dispersal accounts for a larger contribution to gene flow and may maintain high genetic diversity.

Ornamentation, such as the showy plumage of birds, is widespread among female vertebrates, yet the evolutionary pressures shaping female ornamentation remain uncertain. In part this is due to a poor understanding of the mechanistic route to ornamentation in females. To address this issue, we evaluated the evolutionary history of ornament expression in a tropical passerine bird, the White-shouldered Fairywren, whose females, but not males, strongly vary between populations in occurrence of ornamented black-and-white plumage. We first use phylogenomic analysis to demonstrate that female ornamentation is derived and that female ornamentation evolves independently of changes in male plumage. We then use exogenous testosterone in a field experiment to induce partial ornamentation in naturally unornamented females. By sequencing the transcriptome of experimentally induced ornamented and natural feathers, we identify genes expressed during ornament production and evaluate the degree to which female ornamentation in this system is associated with elevated testosterone, as is common in males. We reveal that some ornamentation in females is linked to testosterone and that sexes differ in ornament-linked gene expression. Lastly, using genomic outlier analysis we identify a candidate melanogenesis gene that lies in a region of high genomic divergence among populations that is also differentially expressed in feather follicles of different female plumages. Taken together, these findings are consistent with sex-specific selection favoring the evolution of female ornaments and demonstrate a key role for testosterone in generating population divergence in female ornamentation through gene regulation. More broadly, our work highlights similarities and differences in how ornamentation evolves in the sexes.

Patterns across species of intraspecific phenotypic variation with environment can shed light on the underlying drivers of adaptive evolution. Phenotypic variation within a species along tropical elevational gradients is of particular interest because species with narrow elevational ranges may still experience considerably varied environmental conditions. Here, we examine morphological variation in 27 tropical bird species, spanning 11 families and 3 orders, across a 675 m elevational gradient in Western Ecuador. We analyzed a data set of six morphological variables in 3263 individual birds using multivariate analyses of variance (MANOVAs) and canonical correlation analyses (CCAs). We found that morphology varies significantly with elevation in 8 species, and that spatial segregation by age or sex was apparently not responsible for this result. The phenotypic traits that varied with elevation varied strongly by species. To the best of our knowledge, morphological variation over equally short elevational and horizontal distances across a diverse suite of vertebrate species has not previously been demonstrated.

Understanding the factors that shape the diversity and composition of biotic communities in natural and human-modified landscapes remains a key issue in ecology. Here, we evaluate how functional traits, species diversity and community composition of palm species vary in relation to biogeographic variables and forest age in northwest Ecuador. Functional traits capture essential aspects of species’ ecological tradeoffs and roles within an ecosystem, making them useful in determining the ecological consequences of environmental change, but they have not been used as commonly as more traditional metrics of species diversity and community composition. We inventoried palm communities in 965 10 × 10 m plots arrayed in linear transects placed in forests of varying age. Adult palms in forests of younger regeneration stages were characterized by species with greater maximum stem height, greater maximum stem diameter, and solitary stems. The shift in functional features could indicate that shade tolerant palms are more common in old-growth forest. The shift could also reflect the legacy of leaving canopy palms as remnants in areas that were cleared and then allowed to regrow. Moreover, younger forest age was associated with decreased abundance and altered species composition in both juvenile and adult palms, and decreased species richness in adults. These results highlight the importance of retaining intact, old-growth forest to preserve functional and species diversity and highlight the importance of considering multiple aspects of diversity in studies of vegetation communities.

Although confict is often adaptive and necessary to secure limited resources, it is also frequently a costly endeavor. Signals that reliably communicate competitive ability are commonly employed by animals to reduce confict costs. Both male and female signals have the capacity to serve as honest indicators of competitive ability, but the extent to which this occurs in females has received relatively limited attention. We studied how natural feather coloration and experimentally manipulated plumage ornamentation in female white-shouldered fairywrens (Malurus alboscapulatus moretoni) relates to their aggressive behavior. These fairywrens are a useful study system in that female, but not male coloration varies throughout New Guinea. We tested behavioral responses to simulated rivals both prior to and after plumage manipulation via two distinct behavioral assays: simulated territorial intrusions (where both sexes jointly respond to rival intruders) and mirror image simulation (where females are isolated from their mate). Plumage manipulation treatments had no measurable impact on female aggression during mirror image simulation tests, though aggression did decrease over the course of multiple assays. Similarly, using simulated territorial intrusion assays, we found no diference in female aggression with respect to both natural coloration and manipulation treatment. Finally, our correlative analysis of natural feather color also revealed no detectable relationship. These fndings suggest that female white scapular coloration is unlikely to function as a signal of competitive ability in this tropical species with derived female ornamentation and we discuss possible alternative explanations.

Androgens like testosterone mediate suites of physical and behavioral traits across vertebrates, and circulation varies considerably across and within taxa. However, an understanding of the causal factors of variation in circulating testosterone has proven difficult despite decades of research. According to the challenge hypothesis, agonistic interactions between males immediately prior to the breeding season produce the highest levels of testosterone measured during this period. While many studies have provided support for this hypothesis, most species do not respond to male-male competition by elevating testosterone. As a result, a recent revision of the hypothesis (‘challenge hypothesis 2.0’) places male-female interactions as the primary cause of rapid elevations in testosterone circulation in male vertebrates. Here, we offer a test of both iterations of the challenge hypothesis in a tropical bird species. We first illustrate that male White-shouldered Fairywrens (Malurus alboscapulatus) differ by subspecies in plasma androgen concentrations. Then we use a social network approach to find that males of the subspecies with higher androgens are characterized by greater social interaction scores, including more time aggregating to perform sexual displays. Next, we use a controlled experiment to test whether males respond to simulated territorial intrusion and/or courtship competition contexts by elevating androgens. We found that males elevated androgens during territorial intrusions relative to flushed controls, however, males sampled during courtship competitions had greater plasma androgens both relative to controls and males sampled while defending territories. Ultimately, our results are consistent with challenge hypothesis 2.0, as sexual interactions with extra-pair females were associated with greater elevation of androgens than territorial disputes.

Agonistic conflict is ubiquitous throughout taxa, although the intensity of aggression observed is often highly variable across contexts. For socially monogamous species, a coordinated effort by both pair members can improve the chances of successfully warding off challengers and reinforce pair bonds. However, the intensity of aggression exerted by any one pair member may vary with respect to contextual factors, including the intensity of their mate's aggression. Thus, experimentally exploring how individuals respond to potential rivals via multiple assays with varying social contexts can advance our basic understanding of how aggression varies in socially monogamous systems. We used simulated territorial intrusion and mirror image simulation assays to explore this issue in white-shouldered fairywrens (Malurus alboscapulatus moretoni) of Papua New Guinea. While males tended to be more responsive than females during simulated territorial intrusions, females were more aggressive towards their mirrored reflection than males. Further, individual females that were most aggressive in mirror image simulations were the least aggressive during simulated territorial intrusions, whereas males were inconsistent. These results suggest that female behavioral phenotypes appear to be flexible, relative to context. We discuss how multiple commonly used measurements of aggression might in fact measure different types of responses.

Through frugivory and seed dispersal, vertebrates influence plant demography and forest regeneration. Variation in local habitat surrounding fruiting plants can influence frugivore foraging decisions, thereby creating intraspecific variation in seed dispersal services. However, we have little knowledge of drivers of local variation in frugivory. Here, we investigate factors that may influence frugivore diversity and fruit removal at the level of individual plants. We focus on a common understory palm within a continuous Chocó forest with mixed land-use histories in Ecuador. The density of pioneer tree species in the genus Cecropia around focal palms was negatively related to fruit removal and the diversity of frugivores visiting palms. This may relate to the fact that the presence and abundance of Cecropia species often indicate the existence and severity of past disturbances. Local Cecropia density was also related to an overall shift in the frugivore community that corresponded with an increase in fruit removal by lower-quality seed dispersers (rodents). We also found that the local density of fruiting conspecifics was positively related to frugivore diversity, but not fruit removal. Our results provide information on drivers of intraspecific inequalities in plant populations across tropical forest landscapes. The reduction in fruit removal and frugivore diversity associated with local Cecropia abundance suggests that seed dispersal services can be sensitive to fine-scale variation in habitat structure. Furthermore, because Cecropia are often indicative of past disturbances, this indicates that even small-scale habitat degradation by humans can have lasting effects by creating localized pockets of forest unfavored by frugivores.