Publications

Historically, bird song complexity was thought to evolve primarily through sexual selection on males; yet, in many species, both sexes sing and selection pressure on both sexes may be broader. Previous research suggests competition for mates and resources during short, synchronous breeding seasons leads to more elaborate male songs at high, temperate latitudes. Furthermore, we expect male-female song structure and elaboration to be more similar at lower, tropical latitudes, where longer breeding seasons and year-round territoriality yield similar social selection pressures in both sexes. However, studies seldom take both types of selective pressures and sexes into account. We examined song in both sexes in 15 populations of nine-fairy-wren species (Maluridae), a Southern Hemisphere clade with female song. We compared song elaboration (in both sexes) and sexual song dimorphism to latitude and life-history variables tied to sexual and social selection pressures and sex roles. Our results suggest that song elaboration evolved in part due to sexual competition in males: male songs were longer than female songs in populations with low male survival and less male provisioning. Also, female songs evolved independently of male songs: female songs were slower paced than male songs, although only in less synchronously breeding populations. We also found male and female songs were more similar when parental care was more equal and when male survival was high, which provides strong evidence that sex role similarity correlates with male-female song similarity. Contrary to Northern Hemisphere latitudinal patterns, male and female songs were more similar at higher, temperate latitudes. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts with stronger sexual selection. At the same time, selection pressures associated with sex role similarity appear to favor sex role similarity in song structure.

Conspicuous female signals have recently received substantial scientific attention, but it remains unclear if their evolution is the result of selection acting on females independently of males or if mutual selection facilitates female change. Species that express female, but not male, phenotypic variation among populations represents a useful opportunity to address this knowledge gap. White-shouldered fairywrens (Malurus alboscapulatus) are tropical songbirds with a well-resolved phylogeny where female, but not male, coloration varies allopatrically across subspecies. We explored how four distinct signaling modalities, each putatively associated with increased social selection, are expressed in two populations that vary in competitive pressure on females. Females in a derived subspecies (M. a. moretoni) have evolved more ornamented plumage and have shorter tails (a signal of social dominance) relative to an ancestral subspecies (M. a. lorentzi) with drab females. In response to simulated territorial intrusions broadcasting female song, both sexes of M. a. moretoni are more aggressive and more coordinated with their mates in both movement and vocalizations. Finally, M. a. moretoni songs are more complex than M. a. lorentzi, but song complexity does not vary between sexes in either population. These results suggest that correlated phenotypic shifts in coloration and tail morphology in females as well as song complexity and aggression in both sexes may have occurred in response to changes in the intensity of social selection pressures. This highlights increased competitive pressures in both sexes can facilitate the evolution of complex multimodal signals.

Both abiotic environmental conditions and variation in social environment are known to impact the acquisition of sexual signals. However, the influences of abiotic environmental and social factors are rarely compared to each other. Here we test the relative importance of these factors in determining whether and when male red-backed fairy-wrens, Malurus melanocephalus, moult into ornamented breeding plumage, a known sexual signal. One-year-old male red-backed fairy-wrens vary in whether or not they acquire ornamentation, whereas males age 2 years and older vary in their timing of ornament acquisition. It is unclear whether these processes are determined by the same or different factors, and we examined both events using a combination of long-term breeding records and nonbreeding social networks. We found that 1-year-old males that paired prior to the start of the breeding season were more likely to acquire ornamented plumage, but rainfall did not influence whether 1-year-old males acquired ornamented plumage. Thus, for young individuals, social cues appear to play a larger role than abiotic environmental factors in determining ornament acquisition. For older males, timing of ornamented plumage acquisition was constrained by rainfall, with drier nonbreeding seasons leading to poorer physiological condition and later moult dates. Thus, sexual signal variation in older males appears to be a condition-dependent trait, driven by abiotic environmental and physiological factors rather than social cues. These findings reveal that factors influencing sexual signal expression can vary with age when age classes exhibit different forms of signal variation. Our results suggest that social environment may drive sexual signal variation in young individuals, whereas abiotic environmental variation may drive sexual signal variation in older individuals.

Genetic diversity shapes the evolutionary potential of plant populations. For outcrossing plants, genetic diversity is influenced by effective population size and by dispersal, first of paternal gametes through pollen, and then of paternal and maternal gametes through seeds. Forest loss often reduces genetic diversity, but the degree to which it differentially impacts the paternal and maternal contributions to genetic diversity and the spatial scale at which these impacts are most pronounced are poorly understood. To address these questions, we genotyped 504 seedlings of the animaldispersed palm Oenocarpus bataua collected from 29 widely distributed sites across Ecuador and decomposed the contribution of paternal and maternal gametes to overall genetic diversity. The amount of forest cover at a landscape scale (>10 km radius) had an equally significant positive association with both male and female gametic diversity. In addition, there was a significant positive association between forest cover and effective population size. Stronger fine-scale spatial genetic structure for female versus male gametes was observed at sites with low forest cover, but this did not scale up to differences in male versus female gametic diversity. These findings show that reductions in forest cover at spatial scales much larger than those typically evaluated in ecological studies lead to significant, and equivalent, decreases of diversity in both male and female gametes, and that this association between landscape level forest loss and genetic diversity may be driven directly by reductions in effective population size of O. bataua, rather than by indirect disruptions to local dispersal processes.

Environmental conditions influence ecological processes that shape stream community diversity and abundance. Deforestation has the potential to limit available particulate organic matter and raise stream temperatures. The degree to which tropical stream communities are impacted by these changes is likely to differ between systems, but empirical data from tropical regions are lacking. This lack of baseline data hinders conservation policy as well as efforts to better understand biogeographic and anthropogenic impacts on species’ distributions. To fill this knowledge gap, we surveyed 27 sites in six previously unstudied streams across a gradient of deforestation in northwest Ecuador and assessed the degree to which localized deforestation predicted patterns of community composition of fishes. Using general linear mixed models and AICC we found that neither forest fragmentation nor canopy closure was a significant predictor of species richness and found no difference between the species richness of fragmented and continuous sites. However, forest fragmentation was a strong predictor of abundance, occurring in 31 of 31 general linear mixed models, with higher abundance in fragmented forest than in continuous forest. Of 16 species found, eight occurred at five or more sites and one (Pseudochalceus boehlkei), numbered 200 out of 627 individuals. NMDS and SIMPER analysis suggested that community composition differed between fragmented and continuous sites. P. boehlkei, Pseudopoecilia fria, and Astroblepus cf. fissidens species presented in higher abundances in deforested sites, possibly suggesting a less functionally diverse community. This pattern is consistent with neotropical streams that have experienced partial deforestation but not total degradation of habitat

Acoustic signalling is vital to courtship in many animals, yet the role of female vocalizations is understudied. Here, we combine observational and experimental methods to assess the courtship function of the female chatter call in brown-headed cowbirds, Molothrus ater. While the chatter call is likely multifunctional, it is frequently used in social interactions and overlapping duets with males during the breeding season. Based on a combination of focal- and scan-sampling data from large naturalistic aviaries, we did not find support for the hypothesis that the chatter call elicits male attention or encourages continued courtship. However, we did find evidence that the chatter call plays a role in pair bond formation, as females preferentially chattered in response to songs from pair-bond males in the 2 weeks leading up to the median date of first copulation. Females were less selective in male-directed chatter use after copulations began. We also found support for the hypothesis that chatter is used to signal-jam male songs. Frame-by-frame video analysis revealed that the majority of female chatter calls were tightly time-locked to song, occurring less than 500 ms after male vocal onset. To test the effect of signal jamming on male song potency, we designed a laboratory experiment in which male song playbacks were jammed by various recorded stimuli. Natural chatter calls more effectively reduced female copulatory responses to song than high-pass filtered chatter calls, suggesting that the low frequencies in natural chatter (2-4 kHz) are important for interfering with male song and reducing its potency. Our results suggest that sexual conflict is operating in cowbird courtship, with signal jamming serving as a mechanism by which females guard, resist or select their mates. We also discuss ways in which cowbird vocal interactions may function cooperatively to coordinate reproduction or transition females into breeding condition.

We know little of the proximate mechanisms underlying the expression of signaling traits in female vertebrates. Across males, the expression of sexual and competitive traits, including ornamentation and aggressive behavior, is often mediated by testosterone. In the white-shouldered fairywren (Malurus alboscapulatus) of New Guinea, females of different subspecies differ in the presence or absence of white shoulder patches and melanic plumage, whereas males are uniformly ornamented. Previous work has shown that ornamented females circulate more testosterone and exhibit more territorial aggression than do unornamented females. We investigated the degree to which testosterone regulates the expression of ornamental plumage and territorial behavior by implanting free-living unornamented females with testosterone. Every testosterone-treated female produced a male-like cloacal protuberance, and 15 of 20 replaced experimentally plucked brown with white shoulder patch feathers but did not typically produce melanic plumage characteristic of ornamented females. Testosterone treatment did not elevate territorial behavior prior to the production of the plumage ornament or during the active life of the implant. However, females with experimentally induced ornamentation, but exhausted implants, increased the vocal components of territory defense relative to the pretreatment period and also to testosterone-implanted females that did not produce ornamentation. Our results suggest that testosterone induces partial acquisition of the ornamental female plumage phenotype and that ornament expression, rather than testosterone alone, results in elevations of some territorial behaviors.

Carotenoid pigments produce most red, orange and yellow colours in vertebrates. This coloration can serve as an honest signal of quality that mediates social and mating interactions, but our understanding of the underlying mechanisms that control carotenoid signal production, including how different physiological pathways interact to shape and maintain these signals, remains incomplete. We investigated the role of testosterone in mediating gene expression associated with a red plumage sexual signal in red-backed fairywrens (Malurus melanocephalus). In this species, males within a single population can flexibly produce either red/black nuptial plumage or female-like brown plumage. Combining correlational analyses with a field-based testosterone implant experiment and quantitative polymerase chain reaction, we show that testosterone mediates expression of carotenoid-based plumage in part by regulating expression of CYP2J19, a ketolase gene associated with ketocarotenoid metabolism and pigmentation in birds. This is, to our knowledge, the first time that hormonal regulation of a specific genetic locus has been linked to carotenoid production in a natural context, revealing how endocrine mechanisms produce sexual signals that shape reproductive success.

Tropical forest loss and fragmentation and the associated loss in species diversity are increasing in both magnitude and scope. Much attention has been paid to how attributes of forest fragments, such as area and forest structure, impact the diversity and functional composition of vertebrate communities, while more recent work has begun to consider the importance of landscape-level variables, such as surrounding tree cover. Yet, the relative impacts of these factors on species diversity and functional composition remain unclear, particularly among under-studied taxonomic groups.

Many tropical plant species show wide intra-population variation in reproductive timing, resulting in the protracted presence of flowering and fruiting individuals. Various eco-evolutionary drivers have been proposed as ultimate causes for asynchronous phenology, yet little is known about the proximate factors that control reproductive onset among individuals or that influence the proportion of trees producing new inflorescences within a population. We employed a nine-year phenological record from 178 individuals of the hyperdominant, asynchronously flowering canopy palm, Oenocarpus bataua (Arecaceae)¸ to assess whether resource-related variables influence individual- and population-level flowering phenology. Among individuals, access to sunlight increased rates of inflorescence production, while the presence of resource sinks related to current investment in reproduction—developing infructescences—reduced the probability of producing new inflorescences. At the population level, climate anomalies induced by El Niño Southern Oscillation (ENSO) affected the proportion of the population producing inflorescences through time. Moreover, the effects of ENSO anomalies on flowering patterns depended on the prevalence of developing infructescences in the population, with stronger effects in periods of low developing-infructescence frequency. Taken together, these results suggest that resource-related variables can drive phenological differences among individuals and mediate population-level responses to larger-scale variables, such as climate anomalies. Consequently, a greater focus on the role of resource levels as endogenous cues for reproduction might help explain the frequent aseasonal phenological patterns observed among tropical plants, particularly those showing high intra-population asynchrony.