Publications

Optimal foraging theory states that animals should maximize resource acquisition rates with respect to energy expenditure, which may involve alteration of strategies in response to changes in resource availability and energetic need. However, field-based studies of changes in foraging behavior at fine spatial and temporal scales are rare, particularly among species that feed on highly mobile prey across broad landscapes. To derive information on changes in foraging behavior of breeding brown pelicans (Pelecanus occidentalis) over time, we used GPS telemetry and distribution models of their dominant prey species to relate bird movements to changes in foraging habitat quality in the northern Gulf of Mexico. Over the course of each breeding season, pelican cohorts began by foraging in suboptimal habitats relative to the availability of high-quality patches, but exhibited a marked increase in foraging habitat quality over time that outpaced overall habitat improvement trends across the study site. These findings, which are consistent with adjustment of foraging patch use in response to increased energetic need, highlight the degree to which animal populations can optimize their foraging behaviors in the context of uncertain and dynamic resource availability, and provide an improved understanding of how landscape-level features can impact behavior.

Birds present a stunning diversity of plumage colors that have long fascinated evolutionary ecologists. Although plumage coloration is often linked to sexual selection, it may impact a number of physiological processes, including microbial resistance. At present, the degree to which differences between pigment-based vs. structural plumage coloration may affect the feather microbiota remains unanswered. Using quantitative PCR and DGGE profiling, we investigated feather microbial load, diversity and community structure among two allopatric subspecies of White-shouldered Fairywren, Malurus alboscapulatus that vary in expression of melanin-based vs. structural plumage coloration. We found that microbial load tended to be lower and feather microbial diversity was significantly higher in the plumage of black iridescent males, compared to black matte females and brown individuals. Moreover, black iridescent males had distinct feather microbial communities compared to black matte females and brown individuals. We suggest that distinctive nanostructure properties of iridescent male feathers or different investment in preening influence feather microbiota community composition and load. This study is the first to point to structural plumage coloration as a factor that may significantly regulate feather microbiota. Future work might explore fitness consequences and the role of microorganisms in the evolution of avian sexual dichromatism, with particular reference to iridescence.

Aposematic organisms couple conspicuous warning signals with a secondary defense to deter predators from attacking. Novel signals of aposematic prey are expected to be selected against due to positive frequency-dependent selection. How, then, can novel phenotypes persist after they arise, and why do so many aposematic species exhibit intrapopulation signal variability? Using a polytypic poison frog (Dendrobates tinctorius), we explored the forces of selection on variable aposematic signals using 2 phenotypically distinct (white, yellow) populations. Contrary to expectations, local phenotype was not always better protected compared to novel phenotypes in either population; in the white population, the novel phenotype evoked greater avoidance in natural predators. Despite having a lower quantity of alkaloids, the skin extracts from yellow frogs provoked higher aversive reactions by birds than white frogs in the laboratory, although both populations differed from controls. Similarly, predators learned to avoid the yellow signal faster than the white signal, and generalized their learned avoidance of yellow but not white. We propose that signals that are easily learned and broadly generalized can protect rare, novel signals, and weak warning signals (i.e., signals with poor efficacy and/or poor defense) can persist when gene flow among populations, as in this case, is limited. This provides a mechanism for the persistence of intrapopulation aposematic variation, a likely precursor to polytypism and driver of speciation.

The White-shouldered Fairywren (Malurus alboscapulatus) is a tropical passerine bird distributed across much of New Guinea. White-shouldered Fairywrens are among few species of fairywren with exclusively tropical distributions and differ from better studied congeners in Australia because subspecies vary by female, but not male, coloration and morphology. As with many bird species in New Guinea, basic demographic, social, morphological, and breeding data are limited. From 2011 to 2018 we documented the basic biology of two subspecies representing extremes of the female ornamentation spectrum. Both subspecies form groups having an even operational sex ratio and appear to breed year-round. Extra-pair paternity occurs in the subspecies with female ornamentation; comparable data are lacking for the subspecies having unornamented females, but the greater scaled cloacal protuberance volume of males suggests similar or higher extra-pair paternity rates. Females of the ornamented subspecies are generally larger than those lacking ornamentation, but exhibit reduced tail lengths, which is thought to serve as a signal of social dominance in other fairywrens. After first achieving adult-like plumage, males and ornamented females retain ornamented plumage year-round; however, only males in the subspecies with unornamented females appear to exhibit delayed plumage maturation. Our discussion highlights similarities and differences between White-shouldered Fairywren life histories and those of better studied Australian Malurusspecies; we focus on tropical vs. temperate environments and variable female ornamentation, and we identify priorities for future research.

Many urban areas have elevated soil lead concentrations due to prior large-scale use of lead in products such as paint and automobile gasoline. This presents a potential problem for the growing numbers of wildlife living in urbanized areas as lead exposure is known to affect multiple physiological systems, including the nervous system, in vertebrate species. In humans and laboratory animals, low-level lead exposure is associated with neurological impairment, but less is known about how lead may affect the behavior of urban wildlife. We focused on the Northern Mockingbird Mimus polyglottos, a common, omnivorous North American songbird, to gain insights into how lead may affect the physiology and behavior of urban wildlife. We predicted that birds living in neighborhoods with high soil lead concentrations would (a) exhibit elevated lead concentrations in their blood and feathers, (b) exhibit lower body condition, (c) exhibit less diverse and consistent vocal repertoires, and (d) behave more aggressively during simulated conspecific territorial intrusions compared to birds living in neighborhoods with lower soil lead concentrations. Controlling for other habitat differences, we found that birds from areas of high soil lead had elevated lead concentrations in blood and feathers, but found no differences in body condition or vocal repertoires. However, birds from high lead areas responded more aggressively during simulated intrusions. These findings indicate that sub-lethal lead exposure may be common among wildlife living in urban areas, and that this exposure is associated with increased aggression. Better understanding of the extent of the relationship between lead exposure and aggression and the consequences this could have for survival and reproduction of wild animals are clear priorities for future work in this and other urban ecosystems.

The degree to which foraging individuals are able to appropriately modify their behaviors in response to dynamic environmental conditions and associated resource availability can have important fitness consequences. Despite an increasingly refined understanding of differences in foraging behavior between individuals, we still lack detailed characterizations of within-individual variation over space and time, and what factors may drive this variability. From 2014 to 2017, we used GPS transmitters and accelerometers to document foraging movements by breeding adult Brown Pelicans (Pelecanus occidentalis) in the northern Gulf of Mexico, where the prey landscape is patchy and dynamic at various scales. Assessments of traditional foraging metrics such as trip distance, linearity, or duration did not yield significant relationships between individuals. However, we did observe lower site fidelity and less variation in energy expenditure in birds of higher body condition, despite a population-level trend of increased fidelity as the breeding season progressed. These findings suggest that high-quality individuals are both more variable and more efficient in their foraging behaviors during a period of high energetic demand, consistent with a “rich get richer” scenario in which individuals in better condition are able to invest in more costly behaviors that provide higher returns. This work highlights the importance of considering behavioral variation at multiple scales, with particular reference to within-individual variation, to improve our understanding of foraging ecology in wild populations.

Negative density dependence, where survival decreases as density increases, is a wellestablished driver of species diversity at the community level, but the degree to which a similar process might act on the density or frequency of genotypes within a single plant species to maintain genetic diversity has not been well studied in natural systems. In this study, we determined the maternal genotype of naturally dispersed seeds of the palm Oenocarpus bataua within a tropical forest in northwest Ecuador, tracked the recruitment of each seed, and assessed the role of individual-level genotypic rarity on survival. We demonstrate that negative frequency-dependent selection within this species conferred a survival advantage to rare maternal genotypes and promoted population-level genetic diversity. The strength of the observed rare genotype survival advantage was comparable to the effect of conspecific density regardless of genotype. These findings corroborate an earlier, experimental study and implicate negative frequency-dependent selection of genotypes as an important, but currently underappreciated, determinant of plant recruitment and within-species genetic diversity. Incorporating intraspecific genetic variation into studies and theory of forest dynamics may improve our ability to understand and manage forests, and the processes that maintain their diversity

Habitat loss and fragmentation often reduce gene flow and genetic diversity in plants by disrupting the movement of pollen and seed. However, direct comparisons of the contributions of pollen vs. seed dispersal to genetic variation in fragmented landscapes are lacking. To address this knowledge gap, we partitioned the genetic diversity contributed by male gametes from pollen sources and female gametes from seed sources within established seedlings of the palm Oenocarpus bataua in forest fragments and continuous forest in northwest Ecuador. This approach allowed us to quantify the separate contributions of each of these two dispersal processes to genetic variation. Compared to continuous forest, fragments had stronger spatial genetic structure, especially among female gametes, and reduced effective population sizes. We found that within and among fragments, allelic diversity was lower and genetic structure higher for female gametes than for male gametes. Moreover, female gametic allelic diversity in fragments decreased with decreasing surrounding forest cover, while male gametic allelic diversity did not. These results indicate that limited seed dispersal within and among fragments restricts genetic diversity and strengthens genetic structure in this system. Although pollen movement may also be impacted by habitat loss and fragmentation, it nonetheless serves to promote gene flow and diversity within and among fragments. Pollen and seed dispersal play distinctive roles in determining patterns of genetic variation in fragmented landscapes, and maintaining the integrity of both dispersal processes will be critical to managing and conserving genetic variation in the face of continuing habitat loss and fragmentation in tropical landscapes.

In males, testosterone plays a key role in ornament production and linking ornamentation with reproductive behaviors and other traits to produce an integrated phenotype. Less is known about whether females couple testosterone, ornamentation, and aggressive behaviors to achieve female-specific combinations of traits. Ornamentation in females may be the result of correlated expression with male ornamentation, or female traits could arise as the result of sex-specific selection pressures. Resolving between these alternatives is necessary to understand the degree to which selection acts on female traits. The White-shouldered Fairywren (Malurus alboscapulatus) provides a useful context to address these questions because populations vary in degree of female ornamentation, a derived trait, whereas male ornamentation is constant across both populations. We found that ornamented females have higher levels of circulating testosterone and respond more aggressively to experimental territorial intrusions than do unornamented females. These findings are consistent with the idea that, among female White-shouldered Fairywrens, testosterone may mechanistically link plumage and behavioral traits to produce an integrated competitive phenotype, as has been reported for males of closely related species. In contrast, circulating testosterone in males did not differ significantly between populations. More broadly, our findings are consistent with ongoing selection on the mechanisms underlying female ornaments, likely via social selection.

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (Nem) was associated with higher effective number of pollen sources (Nep), higher effective number of parents (Ne) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (Nem) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.