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The genus Uroleucon, and the related genus Macrosiphoniella, represent a large Tertiary radiation of aphids, with a total of about 300 species distributed throughout the world, primarily on host plant species in the family Asteraceae. A molecular phylogenetic study was conducted to identify major clades within Uroleucon and to address the cladistic validity of current subgeneric categories, the evolution of host plant associations, the age of origin, and intercontinental movements in this genus. The seventeen study species included members of the three major subgenera of Uroleucon, species from Europe and North America, one member of Macrosiphoniella, and two outgroups. Data consisted of DNA sequences for three mitochondrial regions and the nuclear gene EF1alpha, for a total of 4287 sites. Nodes supported strongly in both parsimony and maximum likelihood analyses suggest that: (1) Nearctic Uromelan are a monophyletic group branching near the base of the genus and not related to European Uromelan, (2) the New World subgenus Lambersius is possibly monophyletic but is not a tightly related group and is not closely related to other North American species, and (3) Nearctic members of subgenus Uroleucon are a closely related monophyletic group not allied with Nearctic Uromelan or Lambersius. Instead they represent a separate colonization by an Old World ancestor, as they are nested within a strongly supported clade containing European members of both subgenera Uroleucon and Uromelan. Neither of these subgenera is monophyletic. Molecular clock calculations, based on calibrations of mitochondrial divergences from other insects, suggest that Uroleucon + Macrosiphoniella is a relatively recent radiation, probably no more than 5±10 million years old. Although largely con®ned to Asteraceae, this clade did not radiate in parallel with its host plants. Rather, lateral movement between lineages of Asteraceae must have occurred repeatedly. R Systematic Entomology (1999) 24, 85±93

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Ornamental bill color is postulated to function as a condition‐dependent signal of individual quality in a variety of taxonomically distant bird families. Most red, orange, and yellow bill colors are derived from carotenoid pigments, and carotenoid deposition in ornamentation may trade off with their use as immunostimulants and antioxidants or with other physiological functions. Several studies have found that bill color changes in response to physiological perturbations, but how quickly such changes can occur remains unclear. We tested the hypothesis that carotenoid‐based orange bill color of American goldfinches Spinus tristis responds dynamically to rapid changes in physiological stress and reflects short‐term changes in condition. We captured male and female goldfinches and measured bill color in the field and again under captive conditions several hours later. The following day, the captive birds were injected with either immunostimulatory lipopolysaccharide (LPS) or a control saline and changes in bill color were measured over a five day period. Yellow saturation of the bill decreased within 6.5 h between the field and captivity measures on the first day, presumably in response to capture stress. Over the longer experimental period, bill hue and luminance decreased significantly, whereas saturation significantly increased in both LPS and control groups. Bill hue and luminance decreased significantly more in birds treated with LPS than in control birds. Among LPS treated birds, individuals expressing high bill color at the beginning of the experiment lost more color than ‘low‐color’ birds, but still retained higher color at the end of the experiment, suggesting that birds that invest heavily in bill coloration are able to sustain high costs in the face of a challenge. Bill color change may have resulted from rapid reallocation of carotenoids from ornamentation to immune function. However, the complex shifts in bill color over time suggest that bill color may be influenced by multiple carotenoid compounds and/or changes in blood flow or chemistry in vessels just beneath the translucent keratinized outer layer of the bill. We conclude that bill color is a dynamic, condition‐dependent trait that strategically and reliably signals short‐term fluctuations in physiological condition.

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 woodwarblers (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 nonmutually 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.

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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.

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