Troy G. Murphy

Carotenoid-Based Ornaments of Female and Male American Goldfinches (Spinus tristis) Show Sex-Specific Correlations with Immune Function and Metabolic Rate

Conspicuous ornamentation has been linked to immunological and physiological condition in males of many species. In species where both sexes are ornamented, it is unclear whether the signal content of ornaments differs between males and females. We examined the immunological and physiological correlates of carotenoid-based bill and plumage ornamentation in American goldfinches Spinus tristis, a species in which bright orange bills are sexually monomorphic but yellow plumage is sexually dimorphic during the breeding season. Because bill color is dynamic over short periods while plumage color is static over longer time frames, we tested whether these signals have the potential to provide temporal information about immunity and condition. In both sexes, bill color (but not plumage color) was negatively related to leukocyte differential, a measure of recent stress, while plumage color (but not bill color) was positively related to resting metabolic rate. In females, bill color also positively correlated with immunoglobulin Y, a component of acquired immunity, while plumage color positively predicted natural antibody levels, a component of innate immunity. In males, neither bill color nor plumage color predicted immune function, suggesting that the mechanisms underlying these signals vary with sex. Our results demonstrate that dynamic signals such as bill coloration do not merely reflect the same information provided by static signals but that these two classes of signal provide information about different temporal aspects of phenotypic quality. Furthermore, our results indicate that a signal expressed in both sexes has the potential to provide different information depending on the sex of the bearer.

Sex-Role Reversal in Song? Females Sing More Frequently Than Males in the Streak-Backed Oriole

Abstract Birds in which both sexes produce complex song are more common in the tropics than in the temperate north, where typically only males sing. Yet surprisingly little is known about female song characteristics in most tropical species. Here we present a comparison of female and male singing behaviors in the Streak-backed Oriole (Icterus pustulatus), a tropical songbird in which both sexes perform solo songs. Females sing much more frequently than males and produce songs with similar acoustic complexity. Rates of singing by both sexes were higher during breeding than postbreeding while the rates of most other vocalizations did not change, suggesting that song plays an important role in breeding. To our knowledge, this is the first reported species in which females regularly sing at higher rates than males; however, few studies have examined female song in other sexually monomorphic or weakly dimorphic species, so such patterns might not be unique.

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success, performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.

Lack of Assortative Mating for Tail, Body Size, or Condition in the Elaborate Monomorphic Turquoise-Browed Motmot (Eumomota Superciliosa)

Abstract Elaborate male and female plumage can be maintained by mutual sexual selection and function as a mate-choice or status signal in both sexes. Both male and female Turquoise-browed Motmot (Eumomota superciliosa) have long tails that terminate in widened blue-and-black rackets that appear to hang, unattached, below the body of the bird. I tested whether mutual sexual selection maintains the Turquoise-browed Motmots elaborate tail plumage by testing the prediction that mating occurs in an assortative manner for tail plumage. I also tested whether assortative mating occurs for body size, a potential measure of dominance, and for phenotypic condition, a measure of individual quality. Assortative mating was measured (1) within all pairs in the study population, (2) within newly formed pairs, and (3) within experimentally induced pairs that formed after removal of females from stable pairs. Assortative mating was not found for tail plumage, body size, or phenotypic condition in any of these samples. Therefore, there was no support for the “mutual sexual selection” hypothesis. I discuss the hypothesis that the tail is sexually selected in males only, and that natural selection accounts for the evolutionary maintenance of the elaborate female tail. Ausencia de Apareamiento Asociativo con Respecto a la Cola, el Tamaño Corporal o la Condición en Eumomota superciliosa

Migration and the evolution of sexual dichromatism: evolutionary loss of female coloration with migration among wood-warblers

The mechanisms underlying evolutionary changes in sexual dimorphism have long been of interest to biologists. A striking gradient in sexual dichromatism exists among songbirds in North America, including the wood-warblers (Parulidae): males are generally more colourful than females at northern latitudes, while the sexes are similarly ornamented at lower latitudes. We use phylogenetically controlled comparative analysis to test three non-mutually exclusive hypotheses for the evolution of sexual dichromatism among wood-warblers. The first two hypotheses focus on the loss of female coloration with the evolution of migration, either owing to the costs imposed by visual predators during migration, or owing to the relaxation of selection for female social signalling at higher latitudes. The third hypothesis focuses on whether sexual dichromatism evolved owing to changes in male ornamentation as the strength of sexual selection increases with breeding latitude. To test these hypotheses, we compared sexual dichromatism to three variables: the presence of migration, migration distance, and breeding latitude. We found that the presence of migration and migration distance were both positively correlated with sexual dichromatism, but models including breeding latitude alone were not strongly supported. Ancestral state reconstruction supports the hypothesis that the ancestral wood-warblers were monochromatic, with both colourful males and females. Combined, these results are consistent with the hypotheses that the evolution of migration is associated with the relaxation of selection for social signalling among females and that there are increased predatory costs along longer migratory routes for colourful females. These results suggest that loss of female ornamentation can be a driver of sexual dichromatism and that social or natural selection may be a stronger contributor to variation in dichromatism than sexual selection.

Male Bias in the Song Control System despite Female Bias in Song Rate in Streak-Backed Orioles (Icterus pustulatus)

The song control system is a group of discrete interconnected nuclei found in the brains of all songbirds (suborder Passeri). Previous studies have reported a positive relationship between sex differences in song nucleus volumes and sex differences in song behavior across numerous songbird species, with species exhibiting greater sex differences in behavior also exhibiting greater sex differences in the brain. This body of comparative research, however, has failed to incorporate data from a bird species in which females sing more than males. In this study, we examine song nucleus volumes in both sexes of the streak-backed oriole (Icterus pustulatus), a New World blackbird with a female bias in song rate and similar song complexity between the sexes. Results from this neuroanatomical analysis are contrary to what was to be expected from previous research: despite the female bias in song rate, males have a significantly larger HVC and area X song nucleus volumes. Specifically, male HVC was 75% larger than that of females, and male area X was 64% larger than that of females. There was no significant sex difference in the size of the nucleus robustus arcopallialis. The lack of a positive relationship between song nuclei and singing behavior in these orioles demonstrates that our current understanding of song modulation via the song control system may be overly reliant on basic measures such as total volumes.

Dynamic Status Signal Reflects Outcome of Social Interactions, but Not Energetic Stress

Social defeat induces stress-responses in a wide array of vertebrates and can generate winner-loser effects. Dynamic condition-dependent signaling systems that reflect preparation for subsequent agonistic interactions, and thereby mediate winner-loser effects, should be more sensitive to competitive history than to non-social sources of stress. Bill color of female American goldfinches (Spinus tristus) is a dynamic condition-dependent ornament that functions as a signal of competitive status and mediates intrasexual agonistic social interactions. We tested the “social experience signaling hypothesis” in female goldfinches by (1) manipulating a non-social energetic stressor by experimentally elevating flight costs via wing-clipping in free-ranging birds, and (2) manipulating social experience by staging competitive interactions among captive birds. Bill color change did not differ between wing clipped and non-clipped females, even though stress-response, as measured by the heterophil to lymphocyte (H:L) ratio, increased significantly in clipped females relative to unclipped females. In contrast, winners and losers in the social experience experiment differed significantly in the degree and direction of bill color change following social contests, with bill color increasing in winners and decreasing in losers. These results suggest that dynamic bill color of female American goldfinches signals recent social history, but is less sensitive to some stressors stemming from non-social sources, and thereby suggest that signals can evolve sensitivity to specific types of processes relevant to the context in which they are used.

Same trait, different receiver response: unlike females, male American goldfinches do not signal status with bill colour

In species in which both sexes have similar ornamentation, the ornaments often function as sexual or social signals in both sexes. However, males and females may use ornaments in different signalling contexts. We previously demonstrated that carotenoid-based bill colour of female American goldfinches, Spinus tristis, functions as a signal of status during intrasexual, but not intersexual, competition. Here we test whether male bill colour functions as a competitive status signal during both intra- and intersexual contests. We tested whether focal males and females avoided feeding adjacent to taxidermic male models as a function of the models’ experimentally altered bill colour. We additionally tested whether male bill colour functions as a mate choice signal by presenting females with a choice of two live males with experimentally altered bill colour. In the status signal experiment, neither focal males nor females avoided male models with more colourful bills, as was predicted by the status-signalling hypothesis. These results indicate that male bill coloration does not function as a signal of competitive status and that the signal function of male bill colour does not parallel that of female bill colour. In our mate choice experiment, females showed no preference for male bill colour, suggesting that male bill colour may have some yet untested signalling function or that male bill colour may no longer be under selection. Our findings suggest that selection can lead to different signalling strategies in males and females, even in species that express mutual ornamentation.

Colorful displays signal male quality in a tropical anole lizard

Parasites influence colorful ornaments and their behavioral display in many animal hosts. Because coloration and display behavior are often critical components of communication, variation in these traits may have important implications for individual fitness, yet it remains unclear whether such traits are signals of quality in many taxa. We investigated the association between ectoparasitic mite load and the color and behavioral use of the throat fan (dewlap) by male Anolis brevirostris lizards. We found that heavily parasitized lizards exhibited lower body condition, duller dewlaps, and less frequent dewlap displays than less parasitized individuals. Our results thus suggest that highly parasitized individuals invest less in both ornamental color and behavioral display of that color. Because the two components of the signal simultaneously provide information on male quality, this study provides novel support for the long-standing hypothesis that colorful traits may function as social or sexual signals in reptiles.

Blood parasite infection differentially relates to carotenoid-based plumage and bill color in the American goldfinch

Male and female American goldfinches (Spinus tristis) express condition-dependent carotenoid-based plumage and bill coloration. Plumage color is relatively static, as pigments incorporated into feathers during the spring molt cannot be mobilized thereafter. In contrast, bill color is dynamic, reflecting changes in condition over short time periods. Previous studies have shown that male and female ornaments, though similar in expression, are differentially related to measures of immunocompetence, suggesting that the relationship between ornamentation and parasite infection may differ between the sexes. In this study, we evaluate the relationship between condition-dependent ornamentation (plumage and bill color) and blood parasite infection in male and female American goldfinches. We captured goldfinches after completion of the pre-alternate molt and prior to the onset of nesting and assessed prevalence of Trypanosoma parasites via blood smears. Plumage color strongly predicted trypanosome infection: Birds with more colorful plumage were less likely to present infections. In contrast, we detected no relationship between infection and bill color, which in other studies has been shown to dynamically reflect current condition. Sex did not affect the relationship between infection status and either ornament. Together, these results suggest that physiological pathways linking carotenoid ornamentation and infection may vary even within a single species.