House Finches Research Articles

Carotenoid coloration predicts escape performance in the House Finch (Haemorhous mexicanus)

Carotenoid coloration has been repeatedly shown to serve as a sexually selected signal of individual quality. Across different species, individuals showing brighter carotenoid-based signals have been found to have superior foraging abilities, to recover faster from diseases and, in general, to enjoy a better body condition. Experiments with birds have also shown that carotenoid supplementation can enhance flight performance, allowing birds to take off faster from the ground. Healthy, agile individuals should be better prepared to avoid predators, so it could be expected that individuals displaying brighter carotenoid-based coloration would show a higher escape ability from predator attacks. To test this prediction, we measured the escape ability of male House Finches (Haemorhous mexicanus) from a human with a net in a large aviary and related the escape ability of each individual to its breast coloration. Males with redder feathers showed a higher individual ability to escape than duller individuals. The superior flight performance of redder birds would be an important asset in escape from predators, as well as when foraging or maintaining a territory. In the specific case of the House Finch, the higher escape ability of redder individuals could be the reason for their higher overwinter survival rate.

British Ornithologists' Union Records Committee: 42nd Report (October 2013)

British Ornithologists' Union Records Committee: 42nd Report (October 2013)

Changes in concentrations of circulating heat-shock proteins in House Finches in response to different environmental stressors

Heat-shock proteins (HSP) are molecular chaperones that play key roles in the maintenance of cellular homeostasis under variable environmental conditions. Although HSP are frequently used in studies of wild vertebrates as indicators of stress, no one to date has assessed responses of HSP60, HSP70, and HSP90 in the same species to different environmental stressors. We studied changes in the circulating concentrations of HSP60, HSP70, and HSP90 in wild-caught House Finches (Haemorhous mexicanus) in response to multiple and sequential stressful environments, including high temperatures, transportation, and pathogen exposure. House Finches sampled during a period of low-environmental stress with moderate ambient temperatures had low levels of HSP60 and modest levels of HSP70 and HSP90 compared to birds sampled during a presumably more stressful period with high temperatures. After exposure to high-ambient temperatures, transportation in a vehicle, and exposure to Mycoplasma gallisepticum, captive finches were found to have increasingly higher levels of HSP60. HSP70 tended to rise in response to each stressor, but to drop in the weeks between stress challenges. HSP90 levels increased significantly only in response to pathogen challenge. Our observations suggest that HSP60 and HSP70 are indices of a range of stressors in House Finches, whereas HSP90 primarily reflects health state. Cambios en la concentración de proteínas producidas por estrés, causado por el calor (HSP), en Haemorhous mexicanus como respuesta a diferentes estresantes ambientales Proteínas que se producen por el estrés causado por el calor (HSP) son chaperones moleculares que juegan un papel clave en el mantenimiento de la homeostasis celular bajo diferentes condiciones ambientales. Aunque las HSP son utilizadas con frecuencia en estudios de vertebrados silvestres como indicadores de estrés, hasta el momento nadie ha tratado de determinar la respuesta de HSP60, HSP70 y HSP90, en la misma especie, bajo diferentes condiciones ambientales estresantes. Estudiamos los cambios en las concentraciones de HSP60, HSP70 y HSP90, en el sistema circulatorio de individuos de Haemorhous mexicanus silvestres capturados, como respuesta a múltiples estresantes ambientales, incluyendo altas temperaturas, transportación y exposición a patógenos. Los pinzones muestreados durante un periodo de poco estrés ambiental, con temperaturas ambientales moderadas, mostraron bajos niveles de HSP60, y niveles modestos de HSP70 y HSP90, al ser comparados con aves muestreadas, presumiblemente, durante periodos más estresantes (altas temperaturas). Luego de que las aves cautivas fueran expuestas a altas temperaturas, a transporte en vehículos de motor y a Mycoplasma gallisepticum, se encontraron en estas altos niveles de HSP60. El HSP70 tendió a incrementar en respuesta a cada estresante, pero se redujo en las semanas entre la aplicaciones del estresor. Por su parte el HSP90 solo aumentó, significativamente, en respuesta a los patógenos. Nuestras observaciones sugieren que el HSP60 y HSP70 son índices de una gama de estresantes en individuos del pinzón estudiado, mientras que HSP90 refleja el estado de salud del individuo.

Deposition of pathogenic Mycoplasma gallisepticum onto bird feeders: host pathology is more important than temperature-driven increases in food intake

Although ambient temperature has diverse effects on disease dynamics, few studies have examined how temperature alters pathogen transmission by changing host physiology or behaviour. Here, we test whether reducing ambient temperature alters host foraging, pathology and the potential for fomite transmission of the bacterial pathogen Mycoplasma gallisepticum (MG), which causes seasonal outbreaks of severe conjunctivitis in house finches (Haemorhous mexicanus). We housed finches at temperatures within or below the thermoneutral zone to manipulate food intake by altering energetic requirements of thermoregulation. We predicted that pathogen deposition on bird feeders would increase with temperature-driven increases in food intake and with conjunctival pathology. As expected, housing birds below the thermoneutral zone increased food consumption. Despite this difference, pathogen deposition on feeders did not vary across temperature treatments. However, pathogen deposition increased with conjunctival pathology, independently of temperature and pathogen load, suggesting that MG could enhance its transmission by increasing virulence. Our results suggest that in this system, host physiological responses are more important for transmission potential than temperature-dependent alterations in feeding. Understanding such behavioural and physiological contributions to disease transmission is critical to linking individual responses to climate with population-level disease dynamics.

Mycoplasmosis in wildlife: a review

Mycoplasma, the smallest self-replicating life-forms, are responsible for a variety of diseases in humans, domestic animals, insects, and plants. Although the exact role of these organisms in wildlife is not available, now it is increasingly recognized that mycoplasmosis is an emerging disease in many wildlife populations. As only scattered data regarding these are available, an attempt has been made to assemble the available literature. To our knowledge, this is the first review paper regarding mycoplasmosis in wildlife. The outbreaks of mycoplasmoses such as contagious pleuropneumonia (CCPP), infectious keratoconjunctivitis (IKC), Mycoplasma ovipneumoniae and Mycoplasma capricolum subspp. capricolum infections, and contagious agalactia in wild ruminants; mycoplasmosis in raccoons; upper respiratory tract disease (URTD) and Mycoplasma crocodyli and Mycoplasma alligatoris infections in wild reptiles; and house finch conjunctivitis are showing that the prevalence of mycoplasmosis in wildlife has reached an alarming level, thus raising concerns for the conservation of many endangered species in captivity and free-ranging populations. These also emphasize the urgent need for increased surveillance and control of mycoplasma in wildlife. Some control approaches that can be followed are prevention of the spillover from domestic livestock and birds, proper quarantine of newly received animals along with mycoplasma screening, isolation of affected and suspected animals, screening of mycoplasmoses before the release of captive animals into free-ranging population, etc. At the same time, the previous dogma that mycoplasma have strict host specificity should be reconsidered in the light of new findings.

Do infections lead to higher feather mite loads in birds? A test with mycoplasmal conjunctivitis in House Finches (Haemorhous mexicanus)

Abstract. Current evidence suggests that the health effect of avian feather mites is minimal. However, feather mites can still proliferate without effective preening, such as might occur during sickness. House Finches (Haemorhous mexicanus) are prone to infection by Mycoplasma gallisepticum, symptoms of which include conjunctivitis. Impaired vision and lethargy associated with the disease may limit preening ability. We examined trapping data from a 4-year study of Mycoplasma infection dynamics in House Finches in Atlanta, Georgia, to determine whether birds with conjunctivitis had higher feather mite loads. Abundance of feather mites (Proctophyllodes spp.) and conjunctivitis severity were visually scored on >800 House Finches of known age and sex. Feather mite abundance showed a distinct seasonal pattern: mite loads increased in July and remained high through January. Limiting the analyses to these months, we found that average mite scores were higher in birds with (n = 94) than in those without (n = 275)...

Ketocarotenoid circulation, but not retinal carotenoid accumulation, is linked to eye disease status in a wild songbird

Pathogenic or parasitic infections pose numerous physiological challenges to organisms. Carotenoid pigments have often been used as biomarkers of disease state and impact because they integrate multiple aspects of an individual’s condition and nutritional and health state. Some diseases are known to influence carotenoid uptake from food (e.g. coccidiosis) and carotenoid use (e.g. as antioxidants/immunostimulants in the body, or for sexually attractive coloration), but there is relatively little information in animals about how different types of carotenoids from different tissue sources may be affected by disease. Here we tracked carotenoid accumulation in two body pools (retina and plasma) as a function of disease state in free-ranging house finches (Haemorhous mexicanus). House finches in eastern North America can contract mycoplasmal conjunctivitis (Mycoplasma gallisepticum, or MG), which can progress from eye swelling to eye closure and death. Previous work showed that systemic immune challenges in house finches lower carotenoid levels in retina, where they act as photoprotectors and visual filters. We assessed carotenoid levels during the molt period, a time of year when finches uniquely metabolize ketocarotenoids (e.g. 3-hydroxy-echinenone) for acquisition of sexually selected red plumage coloration, and found that males infected with MG circulated significantly lower levels of 3-hydroxy-echinenone, but no other plasma carotenoid types, than birds exhibiting no MG symptoms. This result uncovers a key biochemical mechanism for the documented detrimental effect of MG on plumage redness in H. mexicanus. In contrast, we failed to find a relationship between MG infection status and retinal carotenoid concentrations. Thus, we reveal differential effects of an infectious eye disease on carotenoid types and tissue pools in a wild songbird. At least compared to retinal sources (which appear somewhat more temporally stable than other body carotenoid pools, even to diseases of the eye evidently), our results point to either a high physiological cost of ketocarotenoid synthesis (as is argued in models of sexually selected carotenoid coloration) or high benefit of using this ketocarotenoid to combat infection.

Multiple host transfers, but only one successful lineage in a continent-spanning emergent pathogen

Emergence of a new disease in a novel host is thought to be a rare outcome following frequent pathogen transfers between host species. However, few opportunities exist to examine whether disease emergence stems from a single successful pathogen transfer, and whether this successful lineage represents only one of several pathogen transfers between hosts. We examined the successful host transfer and subsequent evolution of the bacterial pathogen Mycoplasma gallisepticum, an emergent pathogen of house finches (Haemorhous (formerly Carpodacus) mexicanus). Our principal goals were to assess whether host transfer has been a repeated event between the original poultry hosts and house finches, whether only a single host transfer was ultimately responsible for the emergence of M. gallisepticum in these finches, and whether the spread of the pathogen from east to west across North America has resulted in spatial structuring in the pathogen. Using a phylogeny of M. gallisepticum based on 107 isolates from domestic poultry, house finches and other songbirds, we infer that the bacterium has repeatedly jumped between these two groups of hosts but with only a single lineage of M. gallisepticum persisting and evolving in house finches; bacterial evolution has produced monophyletic eastern and western North American subclades.

Chemosignaling diversity in songbirds: Chromatographic profiling of preen oil volatiles in different species

Large foraging seabirds are known to navigate to food sources using their excellent sense of smell, but much less is known about the use of olfaction by the songbirds (passerine birds). Some evidence of individual recognition based on the bird preen oil volatile organic compound (VOC) compositions, which is the main odor source in birds, have been reported for dark-eyed junco and house finch. In this study we have investigated preen oil VOCs in 16 different songbird species and two other small bird species in order to determine whether the VOC compositions follow phylogenetic and evolutionary relatedness. We have used the stir bar sorptive extraction (SBSE) methodology followed by gas chromatography–mass spectrometry (GC–MS) to determine preen oil VOCs during the long light summer conditions for mostly wild caught birds. Large diversity among the VOC compositions was observed, while some compound classes were found in almost all species. The divergent VOC profiles did not follow the phylogenetic family lines among the bird species. This suggests that songbirds may use VOC odors as a mate recognition cue.

Carotenoid-based plumage coloration reflects feather corticosterone levels in male house finches (Haemorhous mexicanus)

Indicator models of sexual selection predict that exaggerated traits communicate information about sender con- dition or quality to conspecific receivers. Environmental chal- lenges have often been considered as one such condition that could be encoded in an ornamental trait, and there is now extensive evidence showing how different stressors (e.g., nutri- tional, parasitological, and environmental) impact sexual signal elaboration. One of the primary means of assessing stress is by quantifying glucocorticoid (corticosterone or cortisol (CORT)) levels. For many ornaments, CORT impairs trait expression; however, the evidence is limited and mixed for one of the classic honest signals in animals, ornamental carotenoid color- ation. In a model species for studies of carotenoid ornamenta- tion (the house finch, Haemorhous mexicanus), we examined the relationship between male plumage redness and feather CORT levels, which serve as an integrated measure of hormone concentration during feather growth. We measured CORT in both tail (melanin-containing) and breast (carotenoid-contain- ing) feathers and found that CORT levels were not different between body regions, but they were negatively correlated with plumage hue, with redder birds having more CORT in feathers. Despiteopposingtraditionalviews onstressandornamentation, our results actually corroborate three other studies showing positive relationships betwee nc arotenoid coloration and CORT levels. Though the molecular mechanisms underlying such a relationship are still unclear, our results suggest that CORT should not be considered as a simple indicator of indi- vidual quality but rather as a mediator of complex allocation decisions or signals of metabolic activity that could link up with more elaborate expression of ornamental traits.

"Homeostatic Hitchhiking": A Mechanism for the Evolutionary Retention of Complex Adaptations

The complexity of organismal organization channels and accommodates novel genomic and developmental modifications. Here, I extend this perspective to suggest that emergent processes that dominate homeostasis-co-option, re-use, and recombination of accumulated elements-can create configurations and dependencies among these elements that strongly reduce the number of evolutionary steps needed for the evolution of precise novel adaptations. Evolutionary retention and environmental matching of such configurations are further facilitated when they include elements of homeostasis that are responsive to particular environmental cues. I apply this perspective to the study of evolution of sex-biased egg-laying in birds, a phenomenon that combines precision, complexity, context-dependency, and reversibility. I show that homeostatic hitchhiking can overcome the main difficulty in the evolution of this adaptation-the perceived necessity of de novo co-evolution of oogenesis, sex-determination, and order of ovulation in each environmental context-something that would require unrealistic expectations of evolutionary rates and population sizes and is not a desirable outcome for a process that needs to retain substantial environmental sensitivity. First, I explain the rationale behind the homeostatic-hitchhiking hypothesis and outline its predictions specifically for studies of sex-bias in order of egg-laying. Second, I show that a combination of self-regulatory and emergent processes and ubiquitous re-use of conserved growth factors make oogenesis particularly amendable to homeostatic hitchhiking. Third, I review empirical evidence for this mechanism in the rapid evolution of adaptive sex-biased order of egg-laying that accompanied colonization of North America by the house finch (Carpodacus mexicanus).

Effects of carotenoid and vitamin E supplementation on oxidative stress and plumage coloration in house finches (Haemorhous mexicanus)

There has been much recent interest from both applied and basic scientists in the broad series of benefits that animals reap from acquiring high concentrations of dietary antioxidants, such as carotenoids and vitamins (e.g., vitamin E, or tocopherol). Most attention has been paid to separate effects of these compounds on, for example, coloration, health state, development, and vision, but because of possible interactions between these lipid-soluble molecules, we are in need of more studies that co-manipulate these substances and examine their possible synergistic impacts on animal physiology and phenotype. We capitalized on a model avian system (the house finch, Haemorhous mexicanus), where extensive information is available on the fitness roles of carotenoids, to test how variation in carotenoid and/or vitamin E concentrations in the diet impacts body accumulation of these compounds, factors related to oxidative damage (e.g., breast muscle and plasma oxidative-stress susceptibility, plasma nitric-oxide levels), and plumage color development. As in a previous study of ours on carotenoids and health in finches, we employed a 2×2 factorial experimental design on birds in both molting and non-molting conditions, to understand how seasonal shifts in carotenoid use (i.e., pigment incorporation into plumage) might alter the accumulation and roles of carotenoids and vitamins. As expected, lutein supplementation increased the level of circulating carotenoids in both experiments and the color of newly molted plumage. By contrast, vitamin E provisioning did not significantly affect plasma carotenoid levels or plumage coloration in either experiment. Interestingly, carotenoid provisioning decreased circulating vitamin E levels during molt, which suggests either molecular competition between carotenoids and tocopherol at the absorption/transport stages or that vitamin E serves as an antioxidant to offset harmful actions that carotenoids may have at very high concentrations. Finally, in both experiments, we found a reduction in breast-muscle oxidative damage for tocopherol-supplemented birds, which constitutes the first demonstration of a protective effect of vitamin E against oxidative stress in wild birds. Taken together, these findings provide an interesting contrast with our earlier work on season-specific physiological benefits of carotenoids in finches and point to complex associations between indicators of antioxidant and oxidative state in wild-caught animals.

Avian Hosts of West Nile Virus in Arizona

West Nile virus (WNV) causes sporadic outbreaks of human encephalitis in Phoenix, Arizona. To identify amplifying hosts of WNV in the Phoenix area, we blood-sampled resident birds and measured antibody prevalence following an outbreak in the East Valley of metropolitan Phoenix during summer, 2010. House sparrow (Passer domesticus), house finch (Haemorhous mexicanus), great-tailed grackle (Quiscalus mexicanus), and mourning dove (Zenaida macroura) accounted for most WNV infections among locally resident birds. These species roost communally after early summer breeding. In September 2010, Culex vector-avian host contact was 3-fold greater at communal bird roosts compared with control sites, as determined by densities of resting mosquitoes with previous vertebrate contact (i.e., blood-engorged or gravid mosquitoes). Because of the low competence of mourning doves, these were considered weak amplifiers but potentially effective free-ranging sentinels. Highly competent sparrows, finches, and grackles were predicted to be key amplifying hosts for WNV in suburban Phoenix.

Prevalence of Blood Parasites in Eastern Versus Western House Finches: Are Eastern Birds Resistant to Infection?

The rapid spread of the bacterial disease, Mycoplasma gallisepticum (MG), throughout the introduced range of house finches (Carpodacus mexicanus) in eastern North America, compared to its slower spread through the native western range, has puzzled researchers and highlights the need to understand the relative differences in health state of finches from both populations. We conducted a light-microscope survey of hemoparasites in populations of finches from Arizona (within the western range) and from Alabama (within the eastern range), and compared our estimates of prevalence to published reports from house finches sampled in both ranges. Of the 33 Arizona birds examined, we recorded hematozoan infections in 16 (48.5%) individuals, compared to 1 infected Alabama bird out of 30 birds examined (3.3%). Based on independent surveys of seven western North American and five eastern North American populations of house finches the average prevalence of blood parasites in western populations is 38.8% (±17.9 SD), while the average prevalence within the eastern range is only 5.9% (±6.1 SD). The average rate of infection among all songbirds sampled in the east is 34.2% (±4.8 SD). Thus, our surveys of wild birds as well as previously published observations point to eastern house finches having a much lower prevalence of blood parasite infections than their western counterparts. Combined with the fact that eastern finches also tend to have lower rates of avian pox infections than do western birds (based on a literature review), these observations suggest that eastern birds have either strong resistance to these infections or high susceptibility and associated mortality.

Parallel Patterns of Increased Virulence in a Recently Emerged Wildlife Pathogen

Author SummaryA long-standing paradox in the study of infectious diseases is why pathogens evolve to cause harm to the very hosts they depend on to survive and reproduce. Research over several decades suggests that this harm, or virulence, is an inevitable by-product of the pathogen replication needed to maximize the chance that a given pathogen will be transmitted to another host. Here we demonstrate that a recently emerged bacterial pathogen of a North American songbird species has gradually become more virulent during each of two emergence events in different regions of the host range. This evolution of higher virulence appears to have been driven by selection for high rates of pathogen replication, because bacterial isolates that are more virulent in finches also attain the highest loads in infected host tissues. Overall, our results indicate that emerging pathogens can evolve to become more virulent in their hosts, at least in the short term, when an increase in the pathogen's ability to replicate is linked with higher virulence. Our findings have important implications for understanding and predicting the severity of disease caused by emerging pathogens in wildlife, domestic animals, and humans.

Chronic Mycoplasma conjunctivitis in house finches: Host antibody response and M. gallisepticum VlhA expression

Previous studies have shown that house finch field isolates of Mycoplasma gallisepticum (MG) vary in virulence and ability to induce an antibody response. After experimental inoculation, MG causes persistent, severe disease in a subset of individuals. In this study, we further characterized MG infection using five field isolates, with an emphasis on chronically diseased birds. After experimental inoculation of house finches, MG load was measured by quantitative PCR and anti-MG antibody responses were measured by ELISAs. Birds with chronic disease had significantly higher pathogen loads and antibody responses than did birds without chronic disease. Using a monoclonal antibody (MAb86) specific for a variant of the MG VlhA adhesin and immunodominant surface protein, we show that VlhA expression differs among MG isolates in this study, and that in vivo VlhA changes occur in house finches infected with MG. Overall, our results suggest that chronic MG disease has a strong pathogen-mediated component.

House Finch Populations Differ in Early Inflammatory Signaling and Pathogen Tolerance at the Peak ofMycoplasma gallisepticumInfection

Host individuals and populations often vary in their responses to infection, with direct consequences for pathogen spread and evolution. While considerable work has focused on the mechanisms underlying differences in resistance-the ability to kill pathogens-we know little about the mechanisms underlying tolerance-the ability to minimize fitness losses per unit pathogen. Here, we examine patterns and mechanisms of tolerance between two populations of house finches (Haemorhous [formerly Carpodacus] mexicanus) with different histories with the bacterial pathogen Mycoplasma gallisepticum (MG). After infection in a common environment, we assessed two metrics of pathology, mass loss and eye lesion severity, as proxies for fitness. We calculated tolerance using two methods, one based on pathology and pathogen load at the peak of infection (point tolerance) and the other based on the integrals of these metrics over time (range tolerance). Alabama birds, which have a significantly longer history of exposure to MG, showed more pronounced point tolerance than Arizona birds, while range tolerance did not differ between populations. Alabama birds also displayed lower inflammatory cytokine signaling and lower fever early in infection. These results suggest that differences in inflammatory processes, which can significantly damage host tissues, may contribute to variation in tolerance among house finch individuals and populations. Such variation can affect pathogen spread and evolution in ways not predictable by resistance alone and sheds light on the costs and benefits of inflammation in wild animals.

Avian Response to Removal of Feral Sheep on Santa Cruz Island, California

Abstract Feral herbivores can affect birds on islands indirectly, by altering habitat structure via impacts on vegetation. I censused birds in an open shrub/grassland habitat on Santa Cruz Island, California, in 1980 and again in 2008, 24 years after feral sheep were removed, in order to assess avian responses to vegetation changes resulting from sheep removal. Vegetation change was characterized by an increased density of both overstory and low-growing shrubs, along with denser herbaceous vegetation, and these characteristics were associated with substantial changes in bird abundance. Species that prefer dense shrubs, such as Spotted Towhees (Pipilo maculatus) and Orange-crowned Warblers (Oreothlypis celata), increased in abundance, whereas species such as House Finches (Carpodacus mexicanus) and Mourning Doves (Zenaida macroura) that prefer open, disturbed areas decreased. Rufous-crowned Sparrows (Aimophila ruficeps) and Island Scrub-Jays (Aphelocoma insularis), both insular endemic taxa, showed a mixed...

House finches (Carpodacus mexicanus) balance investment in behavioural and immunological defences against pathogens.

Infection with parasites and pathogens is costly for hosts, causing loss of nutritional resources, reproductive potential, tissue integrity and even life. In response, animals have evolved behavioural and immunological strategies to avoid infection by pathogens and infestation by parasites. Scientists generally study these strategies in isolation from each other; however, since these defences entail costs, host individuals should benefit from balancing investment in these strategies, and understanding of infectious disease dynamics would benefit from studying the relationship between them. Here, we show that Carpodacus mexicanus (house finches) avoid sick individuals. Moreover, we show that individuals investing less in behavioural defences invest more in immune defences. Such variation has important implications for the dynamics of pathogen spread through populations, and ultimately the course of epidemics. A deeper understanding of individual- and population-level disease defence strategies will improve our ability to understand, model and predict the outcomes of pathogen spread in wildlife.

Evidence from a House Finch (Haemorhous mexicanus) Spleen Transcriptome for Adaptive Evolution and Biased Gene Conversion in Passerine Birds

Identifying genes influenced by natural selection can provide information about lineage-specific adaptations, and transcriptomes generated by next-generation sequencing are a useful resource for identifying such genes. Here, we utilize a spleen transcriptome for the house finch (Haemorhous mexicanus), an emerging model for sexual selection and disease ecology, together with previously sequenced avian genomes (chicken, turkey, and zebra finch), to investigate lineage-specific adaptations within birds. An analysis of 4,398 orthologous genes revealed a significantly higher ratio of nonsynonymous to synonymous substitutions and significantly higher GC content in passerines than in galliforms, an observation deviating from strictly neutral expectations but consistent with an effect of biased gene conversion on the evolutionary rate in passerines. These data also showed that genes exhibiting signs of positive selection and fast evolution in passerines have functional roles related to fat metabolism, neurodevelopment, and ion binding.