Multimodal models of animal sex in scientific literature: breaking binaries leads to a better understanding of ecology and evolution

‘Sex’ is often used to describe a suite of phenotypic and genotypic traits of an organism related to reproduction. However, not all these traits are necessarily linked, and the rhetorical collapse of variation into a single term elides much of the complexity inherent in reproductive phenotypes. We conducted a meta-analysis of scientific literature to investigate how terminology related sexual phenotypes changes over time. We find that the conflation of gender and sex has increased, and there is a mammalian bias in this conflation. We highlight how a more inclusive and expansive framework for sex can clarify studies of sexual diversity

When females compete and males care: Phenotypic differences in the spotted sandpiper

Ecology can shape variation in mating systems. Spotted sandpipers (Actitis macularius) are migratory shorebirds with a sequentially polyandrous mating system. Females compete for multiple mates, and males care for offspring, including through incubation and chick care. A study of a Midwestern population found that females have larger body mass and feather spots, but we do not know whether this pattern extends to other populations of this species. We studied a California population of spotted sandpipers surrounding Mono Lake. We used PCR to determine genetic sex from blood samples, and we used R to compare morphological traits between females and males. We find sexual dimorphism in several morphological traits, including tarsus, wing, and bill length, as well as body mass, suggesting the Midwestern and California populations have evolved similarly, and both have sexually dimorphic morphology

Behavioural response to song and genetic divergence in two subspecies of white-crowned sparrows (Zonotrichia leucophrys)

© 2016 John Wiley & Sons Ltd Divergence in sexual signals may drive reproductive isolation between lineages, but behavioural barriers can weaken in contact zones. Here, we investigate the role of song as a behavioural and genetic barrier in a contact zone between two subspecies of white-crowned sparrows (Zonotrichia leucophrys). We employed a reduced genomic data set to assess population structure and infer the history underlying divergence, gene flow and hybridization. We also measured divergence in song and tested behavioural responses to song using playback experiments within and outside the contact zone. We found that the subspecies form distinct genetic clusters, and demographic inference supported a model of secondary contact. Song phenotype, particularly length of the first note (a whistle), was a significant predictor of genetic subspecies identity and genetic distance along the hybrid zone, suggesting a close link between song and genetic divergence in this system. Individuals from both parental and admixed localities responded significantly more strongly to their own song than to the other subspecies song, supporting song as a behavioural barrier. Putative parental and admixed individuals were not significantly different in their strength of discrimination between own and other songs; however, individuals from admixed localities tended to discriminate less strongly, and this difference in discrimination strength was explained by song dissimilarity as well as genetic distance. Therefore, we find that song acts as a reproductive isolating mechanism that is potentially weakening in a contact zone between the subspecies. Our findings also support the hypothesis that intraspecific song variation can reduce gene flow between populations

SNP calls for Zonotrichia leucophrys nuttalli x Z. l. pugetensis hybrid zone

Dataset includes 6419 SNP loci for 169 individuals across the Zonotrichia leucophrys nuttalli x Z. l. pugetensis hybrid zone. Sampling information, including individual ID, locality, lat/long, sex, and whether song data is available is also provided. Songs are archived at the Borror Laboratory of Bioacoustics at Ohio State University

Differential introgression of a female competitive trait in a hybrid zone between sex-role reversed species

Mating behavior between recently diverged species in secondary contact can impede or promote reproductive isolation. Traditionally, researchers focus on the importance of female mate choice and male-male competition in maintaining or eroding species barriers. Although female-female competition is widespread, little is known about its role in the speciation process. Here, we investigate a case of interspecific female competition and its influence on patterns of phenotypic and genetic introgression between species. We examine a hybrid zone between sex-role reversed, Neotropical shorebird species, the northern jacana (Jacana spinosa) and wattled jacana (J. jacana), in which female-female competition is a major determinant of reproductive success. Previous work found that females of the more aggressive and larger species, J. spinosa, disproportionately mother hybrid offspring, potentially by monopolizing breeding territories in sympatry with J. jacana. We find a cline shift of female body mass relative to the genetic center of the hybrid zone, consistent with asymmetric introgression of this competitive trait. We suggest that divergence in sexual characteristics between sex-role reversed females can influence patterns of gene flow upon secondary contact, similar to males in systems with more typical sex roles

Data from: Behavioral response to song and genetic divergence in two subspecies of white-crowned sparrows (Zonotrichia leucophrys)

Divergence in sexual signals may drive reproductive isolation between lineages, but behavioral barriers can weaken in contact zones. Here, we investigate the role of song as a behavioral and genetic barrier in a contact zone between two subspecies of white-crowned sparrows (Zonotrichia leucophrys). We employed a reduced genomic dataset to assess population structure and infer the history underlying divergence, gene flow and hybridization. We also measured divergence in song and tested behavioral responses to song using playback experiments within and outside the contact zone. We found that the subspecies form distinct genetic clusters, and demographic inference supported a model of secondary contact. Song phenotype, particularly length of the first note (a whistle), was a significant predictor of genetic subspecies identity and genetic distance along the hybrid zone, suggesting a close link between song and genetic divergence in this system. Individuals from both parental and admixed localities responded significantly more strongly to their own song than to the other subspecies song, supporting song as a behavioral barrier. Putative parental and admixed individuals were not significantly different in their strength of discrimination between own and other songs; however, individuals from admixed localities tended to discriminate less strongly, and this difference in discrimination strength was explained by song dissimilarity as well as genetic distance. Therefore, we find that song acts as a reproductive isolating mechanism that is potentially weakening in a contact zone between the subspecies. Our findings also support the hypothesis that intra-specific song variation can reduce gene flow between populations

Individual sample information

Data file includes information on sample ID, sex, age, date collected, geographic locality, morphological measurements, genomic hybrid index, and species identification

Data from: Differential introgression of a female competitive trait in a hybrid zone between sex-role reversed species

Mating behavior between recently diverged species in secondary contact can impede or promote reproductive isolation. Traditionally, researchers focus on the importance of female mate choice and male-male competition in maintaining or eroding species barriers. Although female-female competition is widespread, little is known about its role in the speciation process. Here, we investigate a case of interspecific female competition and its influence on patterns of phenotypic and genetic introgression between species. We examine a hybrid zone between sex-role reversed, Neotropical shorebird species, the northern jacana (Jacana spinosa) and wattled jacana (J. jacana), in which female-female competition is a major determinant of reproductive success. Previous work found that females of the more aggressive and larger species, J. spinosa, disproportionately mother hybrid offspring, potentially by monopolizing breeding territories in sympatry with J. jacana. We find a cline shift of female body mass relative to the genetic center of the hybrid zone, consistent with asymmetric introgression of this competitive trait. We suggest that divergence in sexual characteristics between sex-role reversed females can influence patterns of gene flow upon secondary contact, similar to males in systems with more typical sex roles

Patterns of Song across Natural and Anthropogenic Soundscapes Suggest That White-Crowned Sparrows Minimize Acoustic Masking and Maximize Signal Content

<div><p>Soundscapes pose both evolutionarily recent and long-standing sources of selection on acoustic communication. We currently know more about the impact of evolutionarily recent human-generated noise on communication than we do about how natural sounds such as pounding surf have shaped communication signals over evolutionary time. Based on signal detection theory, we hypothesized that acoustic phenotypes will vary with both anthropogenic and natural background noise levels and that similar mechanisms of cultural evolution and/or behavioral flexibility may underlie this variation. We studied song characteristics of white-crowned sparrows (<i>Zonotrichia leucophrys nuttalli</i>) across a noise gradient that includes both anthropogenic and natural sources of noise in San Francisco and Marin counties, California, USA. Both anthropogenic and natural soundscapes contain high amplitude low frequency noise (traffic or surf, respectively), so we predicted that birds would produce songs with higher minimum frequencies in areas with higher amplitude background noise to avoid auditory masking. We also anticipated that song minimum frequencies would be higher than the projected lower frequency limit of hearing based on site-specific masking profiles. Background noise was a strong predictor of song minimum frequency, both within a local noise gradient of three urban sites with the same song dialect and cultural evolutionary history, and across the regional noise gradient, which encompasses 11 urban and rural sites, several dialects, and several anthropogenic and natural sources of noise. Among rural sites alone, background noise tended to predict song minimum frequency, indicating that urban sites were not solely responsible for driving the regional pattern. These findings support the hypothesis that songs vary with local and regional soundscapes regardless of the source of noise. Song minimum frequency from five core study sites was also higher than the lower frequency limit of hearing at each site, further supporting the hypothesis that songs vary to transmit through noise in local soundscapes. Minimum frequencies leveled off at noisier sites, suggesting that minimum frequencies are constrained to an upper limit, possibly to retain the information content of wider bandwidths. We found evidence that site noise was a better predictor of song minimum frequency than territory noise in both anthropogenic and natural soundscapes, suggesting that cultural evolution rather than immediate behavioral flexibility is responsible for local song variation. Taken together, these results indicate that soundscapes shape song phenotype across both evolutionarily recent and long-standing soundscapes.</p></div

Map of study sites.

<p>Circles indicate sites within city limits of San Francisco, CA, U.S.A. Triangles indicate sites outside city limits. See text for details on each site.</p