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1.  Invasive ants alter foraging and parental behaviors of a native bird 
Introduced species can exert outsized impacts on native biota through both direct (predation) and indirect (competition) effects. Ants frequently become established in new areas after being transported by humans across traditional biological or geographical barriers, and a prime example of such establishment is the red imported fire ant (Solenopsis invicta). Introduced to North America in the 1930's, red imported fire ants are now firmly established throughout the southeastern United States. Although these invasive predators can dramatically impact native arthropods, their effect on vertebrates through resource competition is essentially unknown. Using a paired experimental design, we compared patterns of foraging and rates of provisioning for breeding eastern bluebirds (Sialia sialis) in unmanipulated (control) territories to those in adjacent (treated) territories where fire ants were experimentally reduced. Bluebirds inhabiting treated territories foraged nearer their nests and provisioned offspring more frequently than bluebirds inhabiting control territories with unmanipulated fire ant levels. Additionally, nestlings from treated territories were in better condition than those from control territories, though these differences were largely confined to early development. The elimination of significant differences in body condition towards the end of the nestling period suggests that bluebird parents in control territories were able to make up the food deficit caused by fire ants, potentially by working harder to adequately provision their offspring. The relationship between fire ant abundance and bluebird behavior hints at the complexity of ecological communities and suggests negative effects of invasive species are not limited to taxa with which they have direct contact.
PMCID: PMC3404743  PMID: 22844172
Invasive species; resource competition; parental care; Sialia sialis; Solenopsis invicta
2.  Sex-biased parental investment is correlated with mate ornamentation in eastern bluebirds 
Animal behaviour  2010;79(3):727-734.
Males typically have greater variance in reproductive success than females, so mothers should benefit by producing sons under favorable conditions. Being paired with a better-than-average mate is one such favorable circumstance. High-quality fathers can improve conditions for their offspring by providing good genes, good resources, or both, so females paired to such males should invest preferentially in sons. Ornamentation has been linked to male quality in many birds and, in support of differential allocation theory, females of several avian species invest more in entire broods when paired to attractive mates. Additionally, the females of some bird species apparently manipulate the primary sex-ratio of their broods in relation to the attractiveness of their mates. However, empirical support for a link between mate ornamentation and preferential feeding of sons (another form of biased investment) is lacking. We tested for correlations between sex-biased parental investment and mate plumage colour in the eastern bluebird (Sialia sialis), a species in which juveniles have sexually dichromatic UV-blue plumage. We found that the proportion of maternal feeding attempts to fledgling sons (versus fledgling daughters) was positively correlated with structurally coloured plumage ornamentation of fathers. Additionally, paternal feeding attempts to sons were correlated with plumage ornamentation of mothers and increased in fathers exhibiting breast plumage characteristics typical of older males. These results provide further support for the idea that parental strategies are influenced by mate attractiveness and provide the first evidence that mate ornamentation can influence parental behavior even after offspring have left the nest.
PMCID: PMC2876345  PMID: 20514141
sex-ratio; parental behavior; resource allocation; ornaments; eastern bluebird; Sialia sialis; honest signal; plumage; colour
4.  Diversity of birds in eastern North America shifts north with global warming 
Ecology and Evolution  2012;2(12):3052-3060.
The distribution of diversity along latitudinal and elevation gradients, and the coupling of this phenomenon with climate, is a pattern long recognized in ecology. Hypothesizing that climate change may have altered this pattern over time, we investigated whether the aggregate of reported northward shifts of bird ranges in North America is now detectable in community-level indices such as richness and diversity. Here, we report that bird diversity in North America increased and shifted northward between 1966 and 2010. This change in the relationship of diversity to the latitudinal gradient is primarily influenced by range expansions of species that winter in the eastern United States as opposed to species which migrate to this area from wintering grounds in the tropics. This increase in diversity and its northward expansion is best explained by an increase in regional prebreeding season temperature over the past 44 years.
PMCID: PMC3539000  PMID: 23301172
Breeding Bird Survey; climate change; diversity; spring temperature; trans-Gulf migrant
5.  A Multi-Year Study of Mosquito Feeding Patterns on Avian Hosts in a Southeastern Focus of Eastern Equine Encephalitis Virus 
Eastern equine encephalitis virus (EEEV) is a mosquito-borne pathogen that cycles in birds but also causes severe disease in humans and horses. We examined patterns of avian host use by vectors of EEEV in Alabama from 2001 to 2009 using blood-meal analysis of field-collected mosquitoes and avian abundance surveys. The northern cardinal (Cardinalis cardinalis) was the only preferred host (fed on significantly more than expected based on abundance) of Culiseta melanura, the enzootic vector of EEEV. Preferred hosts of Culex erraticus, a putative bridge vector of EEEV, were American robin (Turdus migratorius), Carolina chickadee (Poecile carolinensis), barred owl (Strix varia), and northern mockingbird (Mimus polyglottis). Our results provide insight into the relationships between vectors of EEEV and their avian hosts in the Southeast and suggest that the northern cardinal may be important in the ecology of EEEV in this region.
PMCID: PMC3083738  PMID: 21540380
7.  Ultrafast Evolution and Loss of CRISPRs Following a Host Shift in a Novel Wildlife Pathogen, Mycoplasma gallisepticum 
PLoS Genetics  2012;8(2):e1002511.
Measureable rates of genome evolution are well documented in human pathogens but are less well understood in bacterial pathogens in the wild, particularly during and after host switches. Mycoplasma gallisepticum (MG) is a pathogenic bacterium that has evolved predominantly in poultry and recently jumped to wild house finches (Carpodacus mexicanus), a common North American songbird. For the first time we characterize the genome and measure rates of genome evolution in House Finch isolates of MG, as well as in poultry outgroups. Using whole-genome sequences of 12 House Finch isolates across a 13-year serial sample and an additional four newly sequenced poultry strains, we estimate a nucleotide diversity in House Finch isolates of only ∼2% of ancestral poultry strains and a nucleotide substitution rate of 0.8−1.2×10−5 per site per year both in poultry and in House Finches, an exceptionally fast rate rivaling some of the highest estimates reported thus far for bacteria. We also found high diversity and complete turnover of CRISPR arrays in poultry MG strains prior to the switch to the House Finch host, but after the invasion of House Finches there is progressive loss of CRISPR repeat diversity, and recruitment of novel CRISPR repeats ceases. Recent (2007) House Finch MG strains retain only ∼50% of the CRISPR repertoire founding (1994–95) strains and have lost the CRISPR–associated genes required for CRISPR function. Our results suggest that genome evolution in bacterial pathogens of wild birds can be extremely rapid and in this case is accompanied by apparent functional loss of CRISPRs.
Author Summary
Documenting the evolutionary changes occurring in pathogens when they switch hosts is important for understanding mechanisms of adaptation and rates of evolution. We took advantage of a novel host–pathogen system involving a bacterial pathogen (Mycoplasma gallisepticum, or MG) and a songbird host, the House Finch, to study genome-wide changes during a host-shift. Around 1994, biologists noticed that House Finches were contracting conjunctivitis and MG from poultry was discovered to be the cause. The resulting epizootic was one of the best documented for a wildlife species, partly as a result of thousands of citizen science observers. We sequenced the genomes of 12 House Finch MG strains sampled throughout the epizootic, from 1994–2007, as well as four additional putatively ancestral poultry MG strains. Using this serial sample, we estimate a remarkably high rate of substitution, consistent with past implications that mycoplasmas are among the fastest evolving bacteria. We also find that an array of likely phage-derived sequences known as CRISPRs has degraded and ceased to recruit new repeats in the House Finch MG strains, as compared to the poultry strains in which it is diverse and rapidly evolving. This suggests that phage dynamics might be important in the dynamics of MG infection.
PMCID: PMC3276549  PMID: 22346765
8.  Climate change and the decline of a once common bird 
Ecology and Evolution  2012;2(2):370-378.
Climate change is predicted to negatively impact wildlife through a variety of mechanisms including retraction of range. We used data from the North American Breeding Bird Survey and regional and global climate indices to examine the effects of climate change on the breeding distribution of the Rusty Blackbird (Euphagus carolinus), a formerly common species that is rapidly declining. We found that the range of the Rusty Blackbird retracted northward by 143 km since the 1960s and that the probability of local extinction was highest at the southern range margin. Furthermore, we found that the mean breeding latitude of the Rusty Blackbird was significant and positively correlated with the Pacific Decadal Oscillation with a lag of six years. Because the annual distribution of the Rusty Blackbird is affected by annual weather patterns produced by the Pacific Decadal Oscillation, our results support the hypothesis that directional climate change over the past 40 years is contributing to the decline of the Rusty Blackbird. Our study is the first to implicate climate change, acting through range retraction, in a major decline of a formerly common bird species.
PMCID: PMC3298949  PMID: 22423330
Boreal forest; Euphagus carolinus; geographical range; global warming; Odonata; Pacific Decadal Oscillation; population declines; Rusty Blackbird
9.  Estimation of Dispersal Distances of Culex erraticus in a Focus of Eastern Equine Encephalitis Virus in the Southeastern United States 
Journal of medical entomology  2010;47(6):977-986.
Patterns of mosquito dispersal are important for predicting the risk of transmission of mosquito-borne pathogens to vertebrate hosts. We studied dispersal behavior of Culex erraticus (Dyar & Knab), a potentially significant vector of eastern equine encephalitis virus (EEEV) that is often associated with foci of this pathogen in the southeastern United States. Using data on the relative density of resting adult female Cx. erraticus around known emergence sites in Tuskegee National Forest, Alabama, we developed a model for the exponential decay of the relative density of adult mosquitoes with distance from larval habitats through parameterization of dispersal kernels. The mean and 99th percentile of dispersal distance for Cx. erraticus estimated from this model were 0.97 and 3.21 km per gonotrophic cycle, respectively. Parameterized dispersal kernels and estimates of the upper percentiles of dispersal distance of this species can potentially be used to predict EEEV infection risk in areas surrounding the Tuskegee National Forest focus in the event of an EEEV outbreak. The model that we develop for estimating the dispersal distance of Cx. erraticus from collections of adult mosquitoes could be applicable to other mosquito species that emerge from discrete larval sites.
PMCID: PMC3052762  PMID: 21175044
eastern equine encephalitis; insect dispersal; GIS-based modeling; Culex erraticus; larval habitat
10.  Vector–Host Interactions in Avian Nests: Do Mosquitoes Prefer Nestlings over Adults? 
The hypothesis that nestlings are a significant driver of arbovirus transmission and amplification is based upon findings that suggest nestlings are highly susceptible to being fed upon by vector mosquitoes and to viral infection and replication. Several previous studies have suggested that nestlings are preferentially fed upon relative to adults in the nest, and other studies have reported a preference for adults over nestlings. We directly tested the feeding preference of nestling and adult birds in a natural setting, introducing mosquitoes into nesting boxes containing eastern bluebirds (Sialia sialis), collecting blood-fed mosquitoes, and matching the source of mosquito blood meals to individual birds using microsatellite markers. Neither nestlings nor adults were fed upon to an extent significantly greater than would be predicted based upon their relative abundance in the nests, although feeding upon mothers decreased as the age of the nestlings increased.
PMCID: PMC2911192  PMID: 20682889
11.  Invasive Fire Ants Reduce Reproductive Success and Alter the Reproductive Strategies of a Native Vertebrate Insectivore 
PLoS ONE  2011;6(7):e22578.
Introduced organisms can alter ecosystems by disrupting natural ecological relationships. For example, red imported fire ants (Solenopsis invicta) have disrupted native arthropod communities throughout much of their introduced range. By competing for many of the same food resources as insectivorous vertebrates, fire ants also have the potential to disrupt vertebrate communities.
Methodology/Principal Findings
To explore the effects of fire ants on a native insectivorous vertebrate, we compared the reproductive success and strategies of eastern bluebirds (Sialia sialis) inhabiting territories with different abundances of fire ants. We also created experimental dyads of adjacent territories comprised of one territory with artificially reduced fire ant abundance (treated) and one territory that was unmanipulated (control). We found that more bluebird young fledged from treated territories than from adjacent control territories. Fire ant abundance also explained significant variation in two measures of reproductive success across the study population: number of fledglings and hatching success of second clutches. Furthermore, the likelihood of bluebird parents re-nesting in the same territory was negatively influenced by the abundance of foraging fire ants, and parents nesting in territories with experimentally reduced abundances of fire ants produced male-biased broods relative to pairs in adjacent control territories.
Introduced fire ants altered both the reproductive success (number of fledglings, hatching success) and strategies (decision to renest, offspring sex-ratio) of eastern bluebirds. These results illustrate the negative effects that invasive species can have on native biota, including species from taxonomically distant groups.
PMCID: PMC3140521  PMID: 21799904
12.  Feeding decisions of eastern bluebirds are situationally influenced by fledgling plumage color 
Behavioral Ecology  2010;21(3):456-464.
The relative amount of resources that avian parents provide to individual offspring within a brood represents a strategy that can have large effects on reproductive success. We tested whether parental feeding decisions of eastern bluebirds Sialia sialis are influenced by offspring plumage color by presenting pairs of differently colored fledglings side by side and observing how they were provisioned by parents. After a control period, we manipulated blue plumage color so that one sibling in each trial became relatively dark and one became relatively bright. During neither the control nor the experimental periods did either parent consistently feed naturally brighter or experimentally brightened sons more than drab sons. Under specific circumstances, however, both parents directed a higher proportion of their feeding attempts to more brightly colored sons. Paternal feeding attempts to brighter offspring during both the control and experimental periods increased in relation to the brightness of these fledglings relative to their brothers. Maternal feeding decision, on the other hand, were influenced by numerous variables during control and experimental periods including the date of the trial, the difference in mass between fledglings, the feeding behavior of fathers during the trial, the relative investment by fathers during the nestling stage, and the amount of UV chroma in fledgling plumage. Taken together, these results suggest that equal provisioning of offspring is the strategy most commonly adopted by eastern bluebirds but more brightly colored offspring will be fed preferentially when resources for offspring are limited.
PMCID: PMC2854528  PMID: 22476433
color; juvenal plumage; ornaments; parent–offspring interactions; plumage; relative parental investment; Sialia sialis
13.  Host Reproductive Phenology Drives Seasonal Patterns of Host Use in Mosquitoes 
PLoS ONE  2011;6(3):e17681.
Seasonal shifts in host use by mosquitoes from birds to mammals drive the timing and intensity of annual epidemics of mosquito-borne viruses, such as West Nile virus, in North America. The biological mechanism underlying these shifts has been a matter of debate, with hypotheses falling into two camps: (1) the shift is driven by changes in host abundance, or (2) the shift is driven by seasonal changes in the foraging behavior of mosquitoes. Here we explored the idea that seasonal changes in host use by mosquitoes are driven by temporal patterns of host reproduction. We investigated the relationship between seasonal patterns of host use by mosquitoes and host reproductive phenology by examining a seven-year dataset of blood meal identifications from a site in Tuskegee National Forest, Alabama USA and data on reproduction from the most commonly utilized endothermic (white-tailed deer, great blue heron, yellow-crowned night heron) and ectothermic (frogs) hosts. Our analysis revealed that feeding on each host peaked during periods of reproductive activity. Specifically, mosquitoes utilized herons in the spring and early summer, during periods of peak nest occupancy, whereas deer were fed upon most during the late summer and fall, the period corresponding to the peak in births for deer. For frogs, however, feeding on early- and late-season breeders paralleled peaks in male vocalization. We demonstrate for the first time that seasonal patterns of host use by mosquitoes track the reproductive phenology of the hosts. Peaks in relative mosquito feeding on each host during reproductive phases are likely the result of increased tolerance and decreased vigilance to attacking mosquitoes by nestlings and brooding adults (avian hosts), quiescent young (avian and mammalian hosts), and mate-seeking males (frogs).
PMCID: PMC3049777  PMID: 21408172
14.  The Effects of West Nile Virus on the Reproductive Success and Overwinter Survival of Eastern Bluebirds in Alabama 
We tested for negative effects of West Nile virus (WNV) on a breeding population of eastern bluebirds in Alabama by comparing fecundity and reproductive success in years before and after the arrival of WNV and by comparing fecundity, reproductive success, and overwinter survival of seropositive and seronegative individuals within the same population in the same years. We found that female bluebirds were more likely to be seropositive than male bluebirds. Age and individual condition did not affect likelihood of being seropositive. Being seropositive for WNV was not associated with any negative effects on reproduction or survival. However, female fecundity was higher in years after WNV compared to years before the arrival of WNV. The reproductive success of males who tested positive for WNV exposure was higher than that of males that were seronegative. Overall, we found no negative effects on reproduction or survival after exposure to WNV.
PMCID: PMC2883462  PMID: 19589058
Aedes; Arbovirus(es); Birds; Culex; Mosquito(es); Vector-borne
15.  Do adult eastern bluebird Sialia sialis males recognize juvenile-specific traits? 
Animal behaviour  2009;77(5):1267-1272.
Juveniles of many avian species possess spotted or mottled plumage that is distinct from the plumage of adults. Such plumage has typically been assumed to aid in camouflaging vulnerable immature birds. Here, we propose that spotty plumage signals juvenile status, thereby decreasing aggression from territorial adults. We tested this hypothesis by measuring the aggressive responses of adult eastern bluebird males to different combinations of simultaneously presented taxidermic mounts. We found that territorial males attacked adult models significantly more than juvenile models, and that they attacked adult models with orange breasts (typical of adults) more frequently than they attacked adult models with spotty breasts (typical of juveniles). We found no difference in attack rates when models with white breasts (a novel trait) were presented with models possessing spotty breasts. These observations indicate that breast colour is a cue used by territorial adults when identifying conspecific intruders, but that adults do not recognize juvenile-specific plumage as such. Adults respond aggressively only to orange-breasted intruders.
PMCID: PMC2703501  PMID: 20161226
Aggression; Eastern Bluebird; Juvenal Plumage; Honest Signals; Sialia sialis; Territoriality; Juveniles
16.  Developing GIS-based eastern equine encephalitis vector-host models in Tuskegee, Alabama 
A site near Tuskegee, Alabama was examined for vector-host activities of eastern equine encephalomyelitis virus (EEEV). Land cover maps of the study site were created in ArcInfo 9.2® from QuickBird data encompassing visible and near-infrared (NIR) band information (0.45 to 0.72 μm) acquired July 15, 2008. Georeferenced mosquito and bird sampling sites, and their associated land cover attributes from the study site, were overlaid onto the satellite data. SAS 9.1.4® was used to explore univariate statistics and to generate regression models using the field and remote-sampled mosquito and bird data. Regression models indicated that Culex erracticus and Northern Cardinals were the most abundant mosquito and bird species, respectively. Spatial linear prediction models were then generated in Geostatistical Analyst Extension of ArcGIS 9.2®. Additionally, a model of the study site was generated, based on a Digital Elevation Model (DEM), using ArcScene extension of ArcGIS 9.2®.
For total mosquito count data, a first-order trend ordinary kriging process was fitted to the semivariogram at a partial sill of 5.041 km, nugget of 6.325 km, lag size of 7.076 km, and range of 31.43 km, using 12 lags. For total adult Cx. erracticus count, a first-order trend ordinary kriging process was fitted to the semivariogram at a partial sill of 5.764 km, nugget of 6.114 km, lag size of 7.472 km, and range of 32.62 km, using 12 lags. For the total bird count data, a first-order trend ordinary kriging process was fitted to the semivariogram at a partial sill of 4.998 km, nugget of 5.413 km, lag size of 7.549 km and range of 35.27 km, using 12 lags. For the Northern Cardinal count data, a first-order trend ordinary kriging process was fitted to the semivariogram at a partial sill of 6.387 km, nugget of 5.935 km, lag size of 8.549 km and a range of 41.38 km, using 12 lags. Results of the DEM analyses indicated a statistically significant inverse linear relationship between total sampled mosquito data and elevation (R2 = -.4262; p < .0001), with a standard deviation (SD) of 10.46, and total sampled bird data and elevation (R2 = -.5111; p < .0001), with a SD of 22.97. DEM statistics also indicated a significant inverse linear relationship between total sampled Cx. erracticus data and elevation (R2 = -.4711; p < .0001), with a SD of 11.16, and the total sampled Northern Cardinal data and elevation (R2 = -.5831; p < .0001), SD of 11.42.
These data demonstrate that GIS/remote sensing models and spatial statistics can capture space-varying functional relationships between field-sampled mosquito and bird parameters for determining risk for EEEV transmission.
PMCID: PMC2841590  PMID: 20181267
17.  Sex-specific costs of reproduction in Eastern Bluebirds Sialia sialis 
The Ibis  2008;150(1):32-39.
In species with bi-parental care, individuals must partition energy between parental effort and mating effort. Typically, female songbirds invest more than males in reproductive activities such as egg-laying and incubation, but males invest more in secondary sexual traits used in attracting mates. Animals that breed more than once within a season must also allocate time and energy between first and subsequent breeding attempts and between current and future breeding seasons. To investigate strategies of reproductive investment by males and females and the consequences of such strategies, we manipulated the size of broods of Eastern Bluebirds Sialia sialis. Pairs with enlarged first broods were less likely to produce a second clutch or took longer to initiate one than pairs with reduced broods. After rearing enlarged broods, females were less likely than males to survive to the following year. Although plumage coloration is a sexually selected trait in Eastern Bluebirds that is influenced by nutritional stress, we did not detect an effect of brood-size manipulation on female coloration. Past research, however, demonstrates that, in males, plumage colour is negatively affected by increasing brood size. We suggest that there are sex-specific strategies of reproductive investment in Eastern Bluebirds, and that researchers should incorporate measures of residual reproductive value in studies of life-history evolution.
PMCID: PMC2756698  PMID: 19809582
brood manipulation; life-history evolution; multiple clutches; survival; trade-offs
18.  The effect of rearing environment on blue structural coloration of eastern bluebirds (Sialia sialis) 
Behavioral ecology and sociobiology  2007;61(12):1839-1846.
We used a brood-size manipulation to test the effect of rearing environment on structural coloration of feathers grown by eastern bluebird (Sialia sialis) nestlings. Ultraviolet (UV)-blue structural coloration has been shown to be sexually selected in this species. Our experimental design took advantage of the growth of UV-blue wing feathers in nestlings that are retained as part of the first nuptial plumage. We cross-fostered nestlings to create enlarged and reduced broods with the purpose of manipulating parental feeding rates and measured the effect on nestling growth and plumage coloration. Brood size influenced feeding rates to offspring, but the effect varied with season. In general, male nestlings reared in reduced broods were fed more often, weighed more, and displayed brighter structural plumage compared to nestlings reared in enlarged broods. Female nestlings appeared to experience less adverse affects of brood enlargement, and we did not detect an effect of brood-size manipulation on the plumage coloration of female nestlings. Measures of plumage coloration in both males and females, however, were correlated to hatching date and nestling mass during feather development. These data provide empirical evidence that environmental quality can influence the development of the blue structural coloration of feathers and that males may be more sensitive to environmental fluctuations than females.
PMCID: PMC2719904  PMID: 19655039
Structural plumage coloration; Condition-dependent traits; Sexual selection; Parental care; Parental effects
19.  A dynamic transmission model of eastern equine encephalitis virus 
Ecological modelling  2006;192(3-4):425-440.
Eastern equine encephalitis virus (EEEV) is one of several arthropod-borne viruses (arboviruses) endemic to the United States. Interactions between arthropod (mosquito) vectors and avian amplification host populations play a significant role in the dynamics of arboviral transmission. Recent data have suggested the hypothesis that an increased rate of successful feeding on young-of-the-year (YOY) birds might play a role in the dynamics of EEEV transmission. To test this hypothesis, we developed a model to explore the effect of the interactions of the vectors and avian host populations on EEEV transmission. Sensitivity analyses conducted using this model revealed eleven parameters that were capable of disproportionately affecting the predicted level of EEEV infection in the vertebrate reservoir and vector populations. Of these, four parameters were related to the interaction of the vector with young-of-the-year birds. Furthermore, adult birds could not substitute for young-of-the-year in initiating and maintaining a predicted enzootic outbreak of EEEV. Taken together, the model predicted that young-of-the-year birds play a key role in establishing and maintaining enzootic outbreaks of EEEV.
PMCID: PMC1364502  PMID: 16501661
Arbovirus; Avian; Mosquitom; Stella; Alabama
20.  Evolutionary transitions and mechanisms of matte and iridescent plumage coloration in grackles and allies (Icteridae) 
Iridescent structural colour is found in a wide variety of organisms. In birds, the mechanisms that create these colours are diverse, but all are based on ordered arrays of melanin granules within a keratin substrate in barbules. The feathers of the grackles and allies in the family Icteridae range in appearance from matte black to iridescent. In a phylogenetic analysis of this clade, we identified several evolutionary transitions between these colour states. To describe a possible mechanistic explanation for the lability of plumage coloration, we used spectrometry, transmission electron microscopy and thin-film optical modelling of the feathers of 10 icterid species from five genera, including taxa with matte black or iridescent feathers. In matte black species, melanin was densely packed in barbules, while in iridescent species, melanin granules were arranged in ordered layers around the edges of barbules. The structured arrangement of melanin granules in iridescent species created optical interfaces, which are shown by our optical models to be critical for iridescent colour production by coherent scattering. These data imply that rearrangement of melanin granules in barbules is a mechanism for shifts between black and iridescent colours, and that the relative simplicity of this mechanism may explain the lability of plumage colour state within this group.
PMCID: PMC1885357  PMID: 17015306
structural colour; sexual selection; cowbirds; plumage colour; thin-film
21.  An experimental test of the contributions and condition dependence of microstructure and carotenoids in yellow plumage coloration 
A combination of structural and pigmentary components is responsible for many of the colour displays of animals. Despite the ubiquity of this type of coloration, neither the relative contribution of structures and pigments to variation in such colour displays nor the relative effects of extrinsic factors on the structural and pigment-based components of such colour has been determined. Understanding the sources of colour variation is important because structures and pigments may convey different information to conspecifics. In an experiment on captive American goldfinches Carduelis tristis, we manipulated two parameters, carotenoid availability and food availability, known to affect the expression of carotenoid pigments in a full-factorial design. Yellow feathers from these birds were then analysed in two ways. First, we used full-spectrum spectrometry and high-performance liquid chromatography to examine the extent to which variation in white structural colour and total carotenoid content was associated with variation in colour properties of feathers. The carotenoid content of yellow feathers predicted two colour parameters (principal component 1—representing high values of ultraviolet and yellow chroma and low values of violet–blue chroma—and hue). Two different colour parameters (violet–blue and yellow chroma) from white de-pigmented feathers, as well as carotenoid content, predicted reflectance measurements from yellow feathers. Second, we determined the relative effects of our experimental manipulations on white structural colour and yellow colour. Carotenoid availability directly affected yellow colour, while food availability affected it only in combination with carotenoid availability. None of our manipulations had significant effects on the expression of white structural colour. Our results suggest that the contribution of microstructures to variation in the expression of yellow coloration is less than the contribution of carotenoid content, and that carotenoid deposition is more dependent on extrinsic variability than is the production of white structural colour.
PMCID: PMC1639519  PMID: 17015356
American goldfinch; Carduelis tristis; carotenoid pigmentation; diet; honest signalling; sexual selection
22.  Mechanisms of evolutionary change in structural plumage coloration among bluebirds (Sialia spp.) 
Combinations of microstructural and pigmentary components of barbs create the colour displays of feathers. It follows that evolutionary changes in colour displays must reflect changes in the underlying production mechanisms, but rarely have the mechanisms of feather colour evolution been studied. Among bluebirds in the genus Sialia, male rump colour varies among species from dark blue to light blue while breast colour varies from blue to rusty. We use spectrometry, transmission electron microscopy and Fourier analysis to identify the morphology responsible for these divergent colour displays. The morphology of blue rump barbs is similar among the three species, with an outer keratin cortex layer surrounding a medullary ‘spongy layer’ and a basal row of melanin granules. A spongy layer is also present in blue breast barbs of mountain bluebirds Sialia currucoides and in rusty breast barbs of western Sialia mexicana and eastern bluebirds Sialia sialis. In blue barbs melanin is basal to the spongy layer, but is not present in the outer cortex or spongy layer, while in rusty barbs, melanin is present only in the cortex. The placement of melanin in the cortex masks expression of structural blue, creating a rusty display. Such shifts in microstructures and pigments may be widespread mechanisms for the evolutionary changes in the colours of feathers and other reflective structures across colourful organisms.
PMCID: PMC1664640  PMID: 16849249
structural colour; pigmentary colour; Fourier analysis; coherent light scattering
23.  Male eastern bluebirds trade future ornamentation for current reproductive investment 
Biology Letters  2005;1(2):208-211.
Life-history theory proposes that organisms must trade-off investment in current and future reproduction. Production of ornamental display is an important component of reproductive effort that has rarely been considered in tests of allocation trade-offs. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet-blue plumage that is correlated with mate acquisition and male competitive ability. To investigate trade-offs between current reproductive effort and the future expression of a sexually selected ornament, we manipulated the parental effort of males by changing their brood sizes. We found that parents provisioned experimentally enlarged broods more often than reduced broods. As predicted by life-history theory, the change in parental effort had a significant effect on the relative plumage ornamentation of males in the subsequent year: males with reduced broods significantly increased in plumage brightness. Moreover, this change in plumage coloration had a direct effect on the timing of breeding in the following season: males that displayed brighter plumage in the year following the manipulation mated with females that initiated egg laying earlier in the season. These data indicate that male bluebirds must trade-off conserving energy for production of future ornamentation versus expending energy for current reproduction.
PMCID: PMC1626227  PMID: 17148168
sexual selection; life-history evolution; trade-offs; structural plumage; parental effort
24.  Carotenoids need structural colours to shine 
Biology Letters  2005;1(2):121-124.
The bright colours of feathers are among the most striking displays in nature and are frequently used as sexual signals. Feathers can be coloured by pigments or by ordered tissue, and these mechanisms have traditionally been treated as distinct modes of display. Here we show that some yellow plumage colour is created both by reflection of light from white structural tissue and absorption of light by carotenoids. Thus, structural components of feathers contribute substantially to yellow ‘carotenoid’ displays, but the effect of variation in structural components on variation in colour displays is, to our knowledge, unstudied. The presence of structural colour in some carotenoid-based colour displays will have to be considered in studies of colour signalling.
PMCID: PMC1626226  PMID: 17148144
plumage colour; nanostructure; sexual selection; honest signalling
25.  Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour. 
Evidence suggests that structural plumage colour can be an honest signal of individual quality, but the mechanisms responsible for the variation in expression of structural coloration within a species have not been identified. We used full-spectrum spectrometry and transmission electron microscopy to investigate the effect of variation in the nanostructure of the spongy layer on expression of structural ultraviolet (UV)-blue coloration in eastern bluebird (Sialia sialis) feathers. Fourier analysis revealed that feather nanostructure was highly organized but did not accurately predict variation in hue. Within the spongy layer of feather barbs, the number of circular keratin rods significantly predicted UV-violet chroma, whereas the standard error of the diameter of these rods significantly predicted spectral saturation. These observations show that the precision of nanostructural arrangement determines some colour variation in feathers.
PMCID: PMC1691395  PMID: 12965009

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