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Hybrid zones offer a window into the processes and outcomes of evolution, from species formation or fusion to genomic underpinnings of specific traits and isolating mechanisms. Sexual selection is believed to be an important factor in speciation processes, and hybrid zones present special opportunities to probe its impact. The manakins (Aves, Pipridae) are a promising group in which to study the interplay of sexual selection and natural hybridization: they show substantial variation across the family in the strength of sexual selection they experience, they readily hybridize within and between genera, and they appear to have formed hybrid species, a rare event in birds. A hybrid zone between two manakins in the genus Manacus is unusual in that plumage and behavioral traits of one species have introgressed asymmetrically into populations of the second species through positive sexual selection, then apparently stalled at a river barrier. This is one of a handful of documented examples of asymmetric sexual trait introgression with a known selective mechanism. It offers opportunities to examine reproductive isolation, introgression, plumage color evolution, and natural factors enhancing or constraining the effects of sexual selection in real time. Here, we review previous work in this system, propose new hypotheses for observed patterns, and recommend approaches to test them.

Males of many animal species perform specialized courtship behaviours to gain copulations with females. Identifying physiological and anatomical specializations underlying performance of these behaviours helps clarify mechanisms through which sexual selection promotes the evolution of elaborate courtship. Our knowledge about neuromuscular specializations that support elaborate displays is limited to a few model species. In this review, we focus on the physiological control of the courtship of a tropical bird, the golden-collared manakin, which has been the focus of our research for nearly 20 years. Male manakins perform physically elaborate courtship displays that are quick, accurate and powerful. Females seem to choose males based on their motor skills suggesting that neuromuscular specializations possessed by these males are driven by female choice. Male courtship is activated by androgens and androgen receptors are expressed in qualitatively and quantitatively unconventional ways in manakin brain, spinal cord and skeletal muscles. We propose that in some species, females select males based on their neuromuscular capabilities and acquired skills and that elaborate steroid-dependent courtship displays evolve to signal these traits.

Sexual success of males in lek breeding systems is often determined by display activities that require high energetic costs and may represent honest signals of male quality to females. In this context, manakins (Aves, Pipridae) evolved behavioral, morphological, and physiological adaptations that enable complex display maneuvers, and a high rate of fruit ingestion is needed to cope with the energetic costs of lekking activities. We examined the relationship between different displays (rolled-snaps, flights-down, and snap-jumps) and fruit consumption (a proxy for energy intake) by White-bearded Manakin males (Manacus manacus). We conducted a fruit supplementation experiment and recorded male displays in 23 arenas on 5 leks in southeastern Brazil. The flight-down was the display most related to fruit consumption (and energy refueling) by males of the White-bearded Manakin. This display seems to be decisive to the mating success of Manacus males; once inside a lek area, females are driven to display arenas by flights-down of males as an obligatory stage before copulation. This study opens a promising avenue for further field experiments designed to test how local food resources differentially impact display behavior among males in leks.

Manakins are prominent fruit-eaters and seed dispersers in the Neotropics. Most manakin species establish lek areas where males devote long periods of time displaying to attract mates, interspersed with brief absences to feed on fruits located near lek areas. We compared the frequency of visits to plants, fruit handling behavior, and number of fruits ingested by the lek-forming Manacus manacus (White-bearded Manakin) and species of non-lekking birds in two species of fruiting trees (Miconia rigidiuscula and Ocotea pulchella) in lek and non-lek areas during the lek and non-lek periods of M. manacus in a threatened Atlantic forest ecosystem. During the non-lek season, M. manacus and non-lekking birds did not differ in the frequency of visits to Miconia trees near lek and non-lek areas. However, M. manacus swallowed a higher number of fruits near leks than non-lekking birds, while the opposite was true in non-lek areas. During the lek season, M. manacus visited Ocotea trees more frequently and swallowed more fruits than non-lekking birds in lek areas. No birds were recorded on Ocotea trees in non-lek areas. This study provides an example of context dependence in the quantity component of seed dispersal effectiveness in which the lek breeding system of a frugivorous species influences the identity of seed removers and the quantity of seeds removed in and around lek areas. Received 26 November 2012. Accepted 2 February 2013.

Descriptions of lek-breeding in White-bearded Manakins (Manacus manacus) first appeared 55 years ago from studies conducted in tropical Trinidad. No published studies on lek activity, however, exist from subtropical areas of the Neotropics. Herein, annual lek phenology of M. manacus in a subtropical region was analysed and compared with studies conducted in Trinidad. Day-long observations were conducted in nine male territories from three leks. Permanence of lekking males in their territories was positively correlated with day length. When males are more active in the subtropical region, males from Trinidad are less active and vice versa. Additional new information about this manakin's lek activity was disclosed: residents stayed in their courts for up to 7% of the day length, the frequency and duration of display bouts of residents varied throughout the year whereas those of juveniles did not, and females made more visits to courts of residents who displayed for longer bouts.

Understanding the mechanism(s) that favour cooperation among individuals competing for the same resources provides direct insights into the evolution of grouping behaviour. In a hybrid zone between golden- ⁄yellow-collared (Manacus vitellinus) and whitecollared (Manacus candei) manakins, males form aggregations composed of white and yellow males solely to attract females (‘mixed leks’). Previous work shows that yellow males in these mixed leks experience a clear mating advantage over white males, resulting in the preferential introgression of yellow plumage allele(s) into the white species. However, the yellow male mating advantage only occurs in mixed leks with high frequencies of yellow males, and only a few of these males probably mate. Hence, it remains unclear why unsuccessful males join leks. Here, we used microsatellite markers to estimate pairwise relatedness among males within and between leks to test whether indirect genetic benefits of helping kin (‘kin selection’) can promote grouping. We found that yellow males are significantly more related to each other within than between leks, while relatedness among white males did not differ within and between leks. This suggests that yellow males may indirectly enhance their own reproductive success by preferentially lekking with relatives because yellow plumage is under positive frequency-dependent selection (positive FDS). Our results are consistent with the hypothesis that kin selection may promote grouping and facilitate positive FDS for yellow males, mediating the movement of yellow plumage across this hybrid zone.

Hybrid zones are often characterized by narrow, coincident clines for diverse traits, suggesting that little introgression occurs across them. However, this pattern may result from a bias in focussing on traits that are diagnostic of parental populations. Such choice of highly differentiated traits may cause us to overlook differential introgression in nondiagnostic traits and to distort our perception of hybrid zones. We tested this hypothesis in an avian hybrid zone by comparing cline structure in two sets of molecular markers: isozyme and restriction fragment length polymorphism markers chosen for differentiation between parental forms, and microsatellite markers chosen for polymorphism. Two cline-fitting methods showed that cline centre positions of microsatellite alleles were more variable than those of isozyme and restriction fragment length polymorphism markers, and several were significantly shifted from those of the diagnostic markers. Cline widths of microsatellite alleles were also variable and two- to eightfold wider than those of the diagnostic markers. These patterns are consistent with the idea that markers chosen for differentiation are more likely to be under purifying selection, and studies focussed on these markers will underestimate overall introgression across hybrid zones. Our results suggest that neutral and positively selected alleles may introgress freely across many hybrid zones without altering perceived boundaries between hybridizing forms.

Although the power of multi-locus data in estimating species trees is apparent, it is also clear that the analytical methodologies for doing so are still maturing. For example, of the methods currently available for estimating species trees from multiocus data, the Bayesian method introduced by Liu and Pearl (2007; BEST) is the only one that provides nodal support values. Using gene sequences from five nuclear loci, we explored two analytical methods (deep coalescence and BEST) to reconstruct the species tree of the five primary Manacus OTUs: M. aurantiacus, M. candei, M. vitellinus, populations of M. manacus from west of the Andes (M. manacus (w)), and populations of M. manacus from east of the Andes (M. manacus (e)). Both BEST and deep coalescence supported a sister relationship between M. vitellinus and M. manacus (w). A lower probability tree from the BEST analysis and one of the most parsimonious deep coalescence trees also supported a sister relationship between M. candei and M. aurantiacus. Because hybrid zones connect the distributions of most Manacus species, we examined the potential influence of post-divergence gene flow on the sister relationship of parapatrically distributed M. vitellinus and M. manacus (w). An isolation-with-migration (IM) analysis found relatively high levels of gene flow between M. vitellinus and M. manacus (w). Whether the gene flow is obscuring a true sister relationship between M. manacus (w) and M. manacus (e) remained unclear, pointing to the need for more detailed models accommodating multispecies, multilocus DNA sequence data. (Gene trees; incomplete lineage sorting; manakin; phylogeny; species tree.)

Summary The “dancing rings” of Maracus m. trinitatus are almost invariably constructed in small clearings on the forest floor so that whippy saplings grow from their edges to form spring boards as the manakins ricochet back and forth across the rings. Four sounds are made by both sexes, the snap, the “comb” noise, the growling “grrrrt” and the whirring wings. There is no satisfactory explanation of how these sounds are produced. Apart from casual visits, the hens suddenly all appear at about the same time during the morning to dance with the cocks—one hen to a ring. The only exception to this was a pair that spent most of the day together at their ring. Usually at about 9.30 a.m., all the cocks had drifted away, and then nearly every ring was occupied by a hen, dancing back and forth and producing, although not quite as loudly, the same sounds as produced by the cocks. During the period the manakins were under observation (11 February to 16 March), no quarrelling took place among the cocks although their rings were often less than a foot apart; nor were the hens pursued or mated when they danced with the males. This probably indicates that the pairs using the dance rings were already mated or even with nests.

In order to attract females, male golden-collared manakins gather in leks and perform a complex display consisting of acrobatics accompanied by loud "wingsnapping". During this display, males show off their yellow beard and yellow, black, and green plumage that is striking in comparison to the dull green plumage of young males and females. We investigated the role of testosterone (T) in activating the display of manakins and in stimulating the growth of the adult male plumage. T regulates song, copulation, and territorial aggression in temperate species. In tropical species, however, T levels can be relatively low year round, which has raised questions about the involvement of T in courtship display and male aggression in these species. In neither temperate nor tropical species has the role of hormones in the shift from juvenile to adult plumage been well studied. Therefore, we implanted green-plumaged birds and adult males with either a T pellet or an inert pellet (controls) and observed the display behaviors of these birds in the field and in captivity. In captive birds, we also plucked feathers from sexually dimorphic regions and observed color and regeneration rate of new feathers. We found that birds implanted with T increased several display behaviors compared to controls. All plucked feathers grew back the same color as prior to treatment; however, we observed some differences in feather growth rate between T-treated birds and controls.

The evolution of colorful plumage has been dramatic in lekking species. Several studies show that the size of colorful traits influence female choice in leks; however, relatively little is known about the specific function of color, in particular its spectral properties, in lekking taxa. To determine the importance of color in a lekking species, we monitored the mating success of male golden-collared manakins, Manacus vitellinus, and related this to spectral measures of their colorful plumage, as well as other morphological and behavioral traits shown to be important in other lekking species. We found that lek centrality, male body size, and plumage brightness were associated with male mating success. Only plumage brightness, however, entered a multiple regression model, indicating that plumage is the overall best predictor of mating success. These results provide evidence that the spectral properties of colorful plumage predict male mating success in a lekking species and provide important insight into why many lekking birds are dichromatic and elaborate in coloration. Copyright 2006.

How does one species become two? What is the relative importance of selection vs. drift? How many genes are involved, what are their effect sizes and where are they located in the genome? The answers to these questions will inevitably vary from one species to another, but by addressing these questions in a diverse array of taxa, we will gradually piece together a more complete picture of how evolution works. In this issue of Molecular Ecology, Parchman et al. (2013) have approached these questions by studying the genomic basis of differentiation along a hybrid zone between two Central American bird species, the golden-collared (Manacus vitellinus) and white-collared (Manacus candei) manakins (see Fig. 1). Their work presents an interesting window into the evolution of this hybrid zone using novel analysis tools and a nice example of how to approach the study of population genomics in nonmodel organisms. Figure 1. Open Figure Download Powerpoint slide Hybrid (Manacus candei × Manacus vitellinus; left) and white-collared manakin (M. candei; right). Photograph credits: Robb Brumfield (hybrid) and ©D. Tipling/VIREO (M. candei).

The bearded manakins in the genus Manacus are lekking, neotropical passerines. Male plumage colour varies with geographical location and classification is based solely on these plumage patterns. It has recently been suggested that in this group of birds, plumage patterns may be a misleading taxonomic character. In this study we used microsatellite variation in a collection of museum samples to establish the amount of genetic divergence between the previously described bearded manakin species/subspecies. We found substantial genetic substructuring between species/subspecies and that plumage patterns indeed may be a misleading taxonomic character because the presence of yellow in male nuptial plumage is found in most divergent forms. We did not detect a significant isolation by distance relationship although the P-value was close to significance. Physical barriers such as rivers and mountains may affect gene flow and play a role in shaping genetic structure of the genus Manacus. Accordingly, boundaries between species/subspecies often coincide with large rivers, mountains and seas.

SUMMARYBasic kinematic and detailed physical mechanisms of avian, non-vocal sound production are both unknown. Here, for the first time, field-generated high-speed video recordings and acoustic analyses are used to test numerous competing hypotheses of the kinematics underlying sonations, or non-vocal communicative sounds, produced by two genera of Pipridae, Manacus and Pipra (Aves). Eleven behaviorally and acoustically distinct sonations are characterized, five of which fall into a specific acoustic class of relatively loud, brief, broad-frequency sound pulses, or snaps. The hypothesis that one kinematic mechanism of snap production is used within and between birds in general, and manakins specifically, is rejected. Instead, it is verified that three of four competing hypotheses of the kinematic mechanisms used for producing snaps, namely: (1) above-the-back wing-against-wing claps, (2)wing-against-body claps and (3) wing-into-air flicks, are employed between these two clades, and a fourth mechanism, (4) wing-against-tail feather interactions, is discovered. The kinematic mechanisms used to produce snaps are invariable within each identified sonation, despite the fact that a diversity of kinematic mechanisms are used among sonations. The other six sonations described are produced by kinematic mechanisms distinct from those used to create snaps, but are difficult to distinguish from each other and from the kinematics of flight. These results provide the first detailed kinematic information on mechanisms of sonation in birds in general, and the Pipridae specifically. Further, these results provide the first evidence that acoustically similar avian sonations, such as brief, broad frequency snaps, can be produced by diverse kinematic means, both among and within species. The use of high-speed video recordings in the field in a comparative manner documents the diversity of kinematic mechanisms used to sonate, and uncovers a hidden, sexually selected radiation of behavioral and communicative diversity in the Pipridae.

Manakins (Pipridae) are neotropical birds that usually exhibit delayed plumage maturation (DPM). Thus, while plumage of most adult male manakins is brightly conspicuous, subadult males and females are basically dull-olive green. Although sexual dichromatism in some bird species may be evident only through UV reflectance, this phenomenon, known as hidden sexual dichromatism, has not been previously studied in manakins to compare subadult males and females. Within this framework, we carried out spectrophotometric analyses in searching for hidden sexual dichromatism in the white-bearded manakin Manacus manacus, through comparison of UV spectra in females and subadult males in green plumage. Our results revealed UV reflectance in both sexes in green plumage. Moreover, we found UV spectral differences in homologous color patches between sexes, particularly at belly. Since the observed differences may allow intraspecific sex recognition of individuals in green plumage, our results do not support the female-mimicry hypothesis to explain delayed plumage maturation in the white-bearded manakin. Although our findings dismiss the female mimicry hypothesis, we cannot state whether these results support the non-mutually exclusive cryptic and status signaling hypotheses. We propose then, that dull coloration of subadult males may serve both as a cryptic trait and to limit the energetic costs of acquiring the adult plumage before sexual maturity. Meanwhile, differential UV color traits between sexes in green plumage may allow adult males to avoid unnecessary energy expenditures in courtship displays in the presence of males near leks, and to selectively focus their the courtship displays on females. In accordance with the signaling status hypothesis, subadult males can be recognized both as males and subordinates and consequently may practice courtship displays without suffering aggressions by adult males. Our results highlight the importance to include a wider range of spectrophotometric information analyses for testing hypotheses regarding delayed plumage maturation. This article is protected by copyright. All rights reserved.

A previous study of the hybrid zone in western Panama between white-collared (Manacus candei) and golden-collared manakins (M. vitellinus) documented the unidirectional introgression of vitellinus male secondary sexual traits across the zone. Here, we examine the hybrid zone in greater genetic and morphological detail. Statistical comparisons of clines are performed using maximum-likelihood and nonparametric bootstrap methods. Our results demonstrate that an array of six molecular and two morphometric markers agree in cline position and width. Clines for male collar and belly color are similar in width to the first eight clines, but are shifted in position by at least five cline widths. The result is that birds in intervening populations are genetically and morphometrically very like parental candei, but males have the plumage color of parental vitellinus. Neither neutral diffusion nor nonlinearity of color scales appear to be viable explanations for the large cline shifts. Genetic dominance of vitellinus plumage traits is another potential explanation that will require breeding experiments to test. Sexual selection remains a plausible explanation for the observed introgression of vitellinus color traits in these highly dimorphic, polygynous, lek-mating birds. Two other clines, including a nondiagnostic isozyme locus, are similar in position to the main cluster of clines, but are broader in width. Thus, introgression at some loci is greater than that detected with diagnostic markers. Assuming that narrow clines are maintained by selection, variation in cline width indicates that selection is not uniform throughout the genome and that diagnostic markers are under more intense selective pressure. The traditional focus on diagnostic markers in studies of hybrid zones may therefore lead to underestimates of average introgression. This effect may be more pronounced in organisms with low levels of genetic divergence between hybridizing taxa.

We evaluated the effect and effectiveness of antimony potassium tartrate (tartar emetic) on 137 bird species from 29 families and subfamilies from one humid forest site and two mangrove sites in central Panama. Of the 2,656 birds forced to regurgitate, we obtained 1,942 diet samples comprising 12,608 taxonomically identified items. Seventy birds (2.6%) died after administration of the emetic, with some Trochilidae (Phaethornis) and Pipridae (Pipra, Manacus) being especially sensitive to the chemical. No significant differences were found between the number of items regurgitated and that found in stomach contents. However, regurgitations using tartar emetic appeared to be more efficient than analyses of feces to investigate food preference. We also provide details on how to process samples of hummingbirds to permit pollen (flower) identification.

The processes of adaptation and speciation are expected to shape genomic variation within and between diverging species. Here we analyze genomic heterogeneity of genetic differentiation and introgression in a hybrid zone between two bird species (Manacus candei and M. vitellinus) using 59 100 SNPs, a whole genome assembly, and Bayesian models. Measures of genetic differentiation (FST) and introgression (genomic cline center [a ]a nd rate [b]) were highly heterogeneous among loci. We identified thousands of loci with elevated parameter estimates, some of which are likely to be associated with variation in fitness in Manacus populations. To analyze the genomic organization of differentiation and introgression, we mapped SNPs onto a draft assembly of the M. vitellinus genome. Estimates of FST, a ,a ndb were autocorrelated at very short physical distances (< 100 bp), but much less so beyond this. In addition, average statistical associations (linkage disequilibrium) between SNPs were generally low and were not higher in admixed populations than in populations of the parental species. Although they did not occur with a constant probability across the genome, loci with elevated FST, a ,a ndb were not strongly co-localized in the genome. Contrary to verbal models that predict clustering of loci involved in adaptation and isolation in discrete genomic regions, these results are consistent with the hypothesis that genetic regions involved in adaptive divergence and reproductive isolation are scattered throughout the genome. We also found that many loci were characterized by both exceptional genetic differentiation and introgression, consistent with the hypothesis that loci involved in isolation are also often characterized by a history of divergent selection. However, the concordance between isolation and differentiation was only partial, indicating a complex architecture and history of loci involved in isolation.

Theory predicts that traits under positive selection can rapidly cross a hybrid zone in spite of a substantial barrier to neutral gene flow between hybridizing taxa. An avian hybrid zone between Manacus candei (white-collared manakin) and M. vitellinus (golden-collared manakin) is reported here that displays an unusual pattern of noncoincident clines. Male secondary sexual traits of M. vitellinus have spread into populations that are genetically and morphometrically like M. candei. These birds have a lek breeding system in which male mating success is highly skewed, suggesting that sexual selection is driving male sexual traits across the zone.

Phylogenetic relationships among the 14 manakin genera were inferred from DNA sequence data obtained from both mitochondrial and nuclear DNA loci. Phylogenetic analysis resulted in a well-supported hypothesis that corroborates a sister relationship between tyrant-manakins and the ‘‘core” manakins (Antilophia, Chiroxiphia, Corapipo, Dixiphia, Heterocercus, Ilicura, Lepidothrix, Manacus, Masius, Machaeropterus, Pipra, and Xenopipo). Our data strongly support these core manakin genera as a monophyletic group. Consistent with previous work, we find two major clades within the core manakins, although the placement of the genus Xenopipo with regards to these two clades is ambiguous. Generic relationships within these clades are generally well resolved. Although we find some concordance between our study and a previous manakin phylogeny based on syringeal characters, we note several fundamental differences between the phylogenies. Thus, we offer a new phylogenetic hypothesis for Pipridae.

We estimated the annual adult survival (S) of several species of temperate and tropical sexually dimorphic tanagers and manakins from the proportions of first-year and older individuals in museum collections. In the case of the sexually dimorphic species included here, young males through their first year resemble females and can be distinguished from older males. We assumed that the sex ratio among first-year birds is even and estimated the number of immature females (IF) in a sample as equal to the number of immature males (IM), using sex information on museum labels. The number of adult females (AF) is the total number of females (F) minus IF, and survival can be estimated by S = AF/F. We used a Monte Carlo resampling approach to estimate standard errors and 1% and 99% confidence limits on S for samples of size N, based on 1000 sets of N randomly drawn individuals and observed proportions of the F, IM, and AM classes. Estimates of annual survival (S) were 0.60-0.73 in four North American species of Piranga tanagers and 0.50 and 0.62 in two montane Central American species. Values of S in lowland species of Habia and Ramphocelus tanagers ranged between 0.68 and 0.82. Annual survival in Pipra manakins was 0.53-0.68, and values for two Manacus species were 0.69 and 0.70. Where estimates of apparent survival (phi) from capture-recapture studies were available, the two methods produced correlated results with a tendency for phiS. Among tanagers, only Ramphocelus dimidiatus (0.82) and Piranga bidentata (0.50) stood out from the other species, for which S averaged 0.66 +/- 0.05 (SD). Annual survival rates of temperate and tropical species overlapped broadly in this analysis, reinforcing the conclusion that distinctive life-history traits of temperate vs. tropical species cannot be explained completely by the adjustment of reproductive investment in relation to adult survival and the expectation of future life.

Eight microsatellite DNA markers were isolated and characterized from white bearded manakin ( Manacus manacus ) using an enrichment cloning procedure. A large number of alleles (range 9—25), and high levels of observed heterozygosity (mean 0.67) were resolved in 236 individuals. No evidence for linkage disequilibrium or the presence of null alleles was found, indicating that these markers will be useful for examining genetic relatedness, parentage and population structure in manakins.

Hybridization can be an evolutionary creative force by forming new polyploid species, creating novel genetic variation or acting as conduits of potentially advantageous traits between hybridizing forms. Evidence for the latter is often difficult to find because alleles under positive selection can spread rapidly across a hybrid zone and sweep to fixation. In Western Panama, an avian hybrid zone between two species of manakins in the genus Manacus exists where the unidirectional introgression of bright, yellow plumage into a white population provides evidence for the importance of hybrid zones as conduits of advantageous traits. Several lines of indirect evidence suggest that sexual selection favoring yellow plumage drives this asymmetrical spread, but more direct evidence is lacking. Along the edge of the hybrid zone, both yellow- and white-collared manakins are found in the same mating arenas or leks and compete for the same females ("mixed leks"), providing us with a unique opportunity to understand the dynamics of yellow plumage introgression. We studied these mixed leks to determine whether yellow males have a mating advantage over white males and, if so, whether the mating advantage is driven by male-male interactions, female choice, or both. We found that yellow males mated more than white males, suggesting that sexual selection favoring yellow males can, indeed, explain the spread of yellow plumage. However, we found that this advantage occurred only in mixed leks where the frequency of yellow males is greater than white males. This suggests that the advantage of yellow males may depend on the presence of other yellow males, which may slow the rate of introgression in leks where yellow frequency is low such as in areas where yellow males are beginning to colonize the white population. This, along with the geographic barrier posed by major rivers in the hybrid zone, may initially limit or slow the spread of yellow plumage. Finally, we found that yellow and white males were similar in aggression and body size, and held comparable positions within leks. Because these traits or factors are often important in or dictated by aggressive male-male interactions, these comparisons indicate that male-male interaction is not the primary mechanism for the spread of yellow plumage. However, white and yellow males received similar numbers of courtship visits from females but differed in the number of matings, suggesting that females actively rejected white in favor of yellow males. Our results indicate that sexual selection by female choice has driven the unidirectional introgression of yellow plumage into the white population, providing a mechanism for how hybrid zones act as conduits of novel and advantageous traits.

The piprid genus Manacus is composed of four allospecies that are readily distinguishable by differences in male plumage color. Electrophoretic data for two populations of each of the four forms plus seven outgroup piprid taxa were collected from 32 isozyme loci and used to infer phylogenetic relationships. Each Manacus form was monophyletic, with the exception of M. manacus, in which the trans-Andean (west of the Andes) population was sister to M. vitellinus, rather than to its conspecific cis-Andean (east of the Andes) population. This controversial relationship, supported by the synapomorphic allele PGM-2b as well as allele frequencies at ADA, GOT-1 and LGG, is consistent with general biogeographic patterns in the region, but indicates that male plumage color is an unreliable taxonomic marker. Reconstruction of male plumage color on the tree confirms that gold plumage is a derived state in M. vitellinus, a finding consistent with the possibility that gold plumage is an evolutionary novelty in ...

I examined the digestive physiology of two avian frugivores, the golden-collared manakin, Manacus vitellinus, and the red-capped manakin, Pipra mentalis, to discover how these birds extract energy from fruit. Using 14 species of fruit in the natural diet of manakins, I examined the assimilation of nutrients from fruit pulp, fruit passage rates, seed passage rates, and gut morphology. Fruits in the manakins' diets had high water content (average, 84%) and low nutrient concentrations (3 kJ/g wet pulp; 17 kJ/g dry pulp; 1% nitrogen/g dry pulp). Manacus and Pipra did not differ in the average assimilation of energy in fruit pulp (63%), although it varied from 37 to 84% depending on fruit species. Assimilation of total nonstructural carbohydrates in the fruit pulp was very high (86–98%) in both species. Gut evacuation was rapid; maximum transit time of a labeled fruit was 30 min. Seeds passed through the gut faster (Manacus: 15 min; Pipra: 12 min) than the accompanying fruit epidermis (both spp: 22 min). Manakins regurgitated large seeds (>5 mm diameter) in 7 to 9 min. Rapid gut passage time, high assimilation of nonstructural carbohydrates, and the selective regurgitation and rapid elimination of bulky seeds enable manakins to process a large volume of food per day. By increasing rates of fruit intake and gut passage, manakins can effectively increase total nutrient uptake. These adaptations of manakins are requisite for harvesting sufficient nutrients from fruit, due to its low nutrient density, high water content, and bulky seeds.