The evolution of immune defense and song complexity in birds

Abstract There are three main hypotheses that explain how the evolution of parasite virulence could be linked to the evolution of secondary sexual traits, such as bird song. First, as Hamilton and Zuk proposed a role for parasites in sexual selection, female preference for healthy males in heavily parasitized species may result in extravagant trait expression. Second, a reverse causal mechanism may act, if sexual selection affects the coevolutionary dynamics of host-parasite interactions per se by selecting for increased virulence. Third, the immuno-suppressive effects of ornamentation by testosterone or limited resources may lead to increased susceptibility to parasites in species with elaborate songs. Assuming a coevolutionary relationship between parasite virulence and host investment in immune defense we used measures of immune function and song complexity to test these hypotheses in a comparative study of passerine birds. Under the first two hypotheses we predicted avian song complexity to be positively related to immune defense among species, whereas this relationship was expected to be negative if immuno-suppression was at work. We found that adult T-cell mediated immune response and the relative size of the bursa of Fabricius were independently positively correlated with a measure of song complexity, even when potentially confounding variables were held constant. Nestling T-cell response was not related to song complexity, probably reflecting age-dependent selective pressures on host immune defense. Our results are consistent with the hypotheses that predict a positive relationship between song complexity and immune function, thus indicating a role for parasites in sexual selection. Different components of the immune system may have been independently involved in this process.     

Pollination by passerine birds: why are the nectars so dilute?

Bird-pollinated flowers are known to secrete relatively dilute nectars (with concentrations averaging 20–25% w/w). Many southern African plants that are pollinated by passerine birds produce nectars with little or no sucrose. Moreover, these hexose nectars are extremely dilute (10–15%). This suggests a link between sugar composition and nectar concentration. Nectar originates from sucrose-rich phloem sap, and the proportion of monosaccharides depends on the presence and activity of invertase in the nectary. Hydrolysis of sucrose increases nectar osmolality and the resulting water influx can potentially convert a 30% sucrose nectar into a 20% hexose nectar, with a 1.56 times increase in volume. Hydrolysis may also increase the gradient for sucrose transport and thus the rate of sugar secretion. When sucrose content and refractometer data were compared, some significant correlations were seen, but the occurrence of sucrose-rich or hexose-rich nectars can also be explained on phylogenetic grounds (e.g. Erythrina and Protea). Hexose nectars may be abundant enough to drip from open flowers, but evaporation leads to much variability in nectar concentration and increases the choices available to pollinators.     

Comparative analysis of avian BMAL1 and CLOCK protein sequences: a search for features associated with owl nocturnal behaviour

Animals differ widely in the phasing of their daily rhythms with respect to daily environmental rhythms. While birds are predominantly day-active, nocturnal activity is a characteristic feature of the order Strigiformes (owls). To study the evolution of owl night-activity cDNA sequences encoding the circadian core oscillator (CCO) proteins BMAL1 and CLOCK were obtained from barn owl (Tyto alba). The predicted proteins showed high sequence identity with their Galliform homologues (BMAL1: 99%; CLOCK: 95.6%). A computer-predicted chicken BMAL1 casein kinase-1 phosphorylation site is absent from T. alba BMAL1, but also absent from homologues of other six bird species (5 orders) (night-active (n=2), day-active (n=4)) indicating no evolutionary association with night activity. Sequence differences between T. alba and Galliform CLOCK frequently involved serine and threonine residues suggesting potential differences in their phosphorylation. The length of a poly-glutamine string in the CLOCK C-terminus varied between and within 25 species (6 orders) examined, however, no discernible feature distinguishing day and night active species was found. No differences were found between day (n=5) and night (n=7)-active species (12 species, 6 orders) in a region of the PER2 protein implicated in altered rhythm phasing in humans. In conclusion the avian CCO components examined showed strong evolutionary conservation. Molecular evolution associated with owl night-activity may have involved alterations in the CCO relationship with ‘output’ genes rather than in the molecular structure of the CCO itself.     

Local survival after fire in Mediterranean shrublands: combining capture-recapture data over several bird species

Post-disturbance survival is a key factor in the onset of secondary successions. Here we analyse capture-recapture data from two before/after disturbance studies to estimate the effect of fire on local bird survival. Analyses of six bird species at two Mediterranean shrubland sites were combined using a meta-analysis approach. Two warblers, Sylvia undata and S. melanocephala, were studied at one site altered by prescribed burning, and five passerines (Luscinia megarhynchos, Turdus merula, Parus major, P. caeruleus and S. melanocephala) at one site disturbed by wildfire. Based on the combined analysis, annual survival probability significantly decreased from 0.49 to 0.18 (i.e. a 64% decline) after the fire. Our results further suggest a trend for a higher decrease in annual survival associated with wildfire (–72%, from 0.51 to 0.14) than with prescribed burning (–35%, from 0.41 to 0.27), although this should be properly tested with a specific experimental design. In S. undata, a decline in survival in the long-term cannot account for the drop in density observed the first spring after fire. We suggest that a decrease in recruitment rate and an increase in the proportion of non-breeders immediately after the fire may contribute more strongly to the decline in the breeding population. Our results tend to support the idea that bird populations may respond to moderate disturbances with noticeable time lags, because of individual site tenacity.     

Behavioral Components of Sex Role Reversal in the Tettigoniid Bushcricket Ephippiger ephippiger

Facultative sex role reversal is found in species of tettigoniid bushcrickets in which males invest heavily in matings or offspring by producing large, nutritious spermatophores. On high-quality diets such species show conventional sex roles, but under low-quality diets males become the choosier sex. Comparative work suggests that Ephippiger ephippiger (Tettigoniidae, Ephippigerinae) has one of the largest described spermatophores (up to 40% of the male body weight). Here we examine the behavior of this species under variable diet conditions in the laboratory and find evidence of sex role reversal under poor-quality diet conditions. We also examine the behavioral components of sex role reversal. In the reversed condition, rejections of mating attempts are almost solely by the male and contests are almost solely between females. Role-reversed males sing less frequently and at a much reduced intensity. We use geographic variation in the calling song of this species to assess the strength of female phonotactic discrimination between races. This is not significantly reduced despite sex role reversal. We therefore suggest that the male and female aspects of the acoustic sexual communication system differ in that male components change most during facultative sex role reversal.     

西双版纳部分地区鸟类多样性初步考察

１９９２年１２月,１９９３年１１月及１９９４年４～５月,对西双版纳自然保护区内外的数个工作点进行了鸟类物种多样性的调查,共记录鸟类１９０种。分析表明雀形目鸟类物种多样性状态尚佳,而非雀形目鸟类类群（科种）多样性及物种多样性状态均恶化趋势,保护区内鸟类物种多样性状态明显较好,傣族墓葬地之龙山因依傍村落,农田及其他多种生境而具有相当丰富的鸟类物种多样性,是保护鸟类物种多样性不可忽视的一个方面,研究表明     

Seasonal shifts in sex ratios of fledgling American kestrels (Falco sparverius paulus): The Early Bird Hypothesis

We document a seasonal shift in the sex ratios of broods produced by resident southeastern American kestrels (Falco sparverius paulus) breeding in nest boxes in Florida. Early in the breeding season, most biased broods were biased towards males, whereas later in the season, most biased broods were biased towards females. Computer-simulated broods subjected to sex-biased egg and/or nestling mortality demonstrate that it is possible that differential mortality produced the pattern of bias that we observed. However, these simulations do not exclude the possibility that female kestrels were manipulating the primary sex ratio of the broods. We present evidence that this sex ratio shift is adaptive: for males we detected breeding as yearlings, all had fledged early the previous season. No such relationship between season and the probability of breeding as a yearling was found for females. We propose the Early Bird Hypothesis as the ecological basis for the advantage of fledg ing early in males. We hypothesize that pre-emptive competition among post-fledging, dispersing males for breeding sites confers an advantage to males fledged early in the season. This hypothesis may explain why a non-migratory population of the Eurasian kestrel (F. tinnunculus) and non-migratory American kestrels breeding in Florida (F. s. paulus) exhibit this seasonal shift in sex ratios, whereas migratory American kestrels (F. s. sparverius) breeding in Saskatchewan, Canada, do not. We discuss the relevance of the Early Bird Hypothesis for other animal species.     

Bird diversity: a predictable function of satellite-derived estimates of seasonal variation in canopy light absorbance across the United States

Aim  To investigate the relationships between bird species richness derived from the North American Breeding Bird Survey and estimates of the average, minimum, and the seasonal variation in canopy light absorbance (the fraction of absorbed photosynthetically active radiation, fPAR) derived from NASA's Moderate Resolution Imaging Spectroradiometer (MODIS).
Location  Continental USA.
Methods  We describe and apply a 'dynamic habitat index' (DHI), which incorporates three components based on monthly measures of canopy light absorbance through the year. The three components are the annual sum, the minimum, and the seasonal variation in monthly fPAR, acquired at a spatial resolution of 1 km, over a 6-year period (2000–05). The capacity of these three DHI components to predict bird species richness across 84 defined ecoregions was assessed using regression models.
Results  Total bird species richness showed the highest correlation with the composite DHI [ R ² = 0.88, P  < 0.001, standard error of estimate (SE) = 8 species], followed by canopy nesters ( R ² = 0.79, P  < 0.001, SE = 3 species) and grassland species ( R ² = 0.74, P  < 0.001, SE = 1 species). Overall, the seasonal variation in fPAR, compared with the annual average fPAR, and its spatial variation across the landscape, were the components that accounted for most ( R ² = 0.55–0.88) of the observed variation in bird species richness.
Main conclusions  The strong relationship between the DHI and observed avian biodiversity suggests that seasonal and interannual variation in remotely sensed fPAR can provide an effective tool for predicting patterns of avian species richness at regional and broader scales, across the conterminous USA.     

A blueprint for vocal learning: auditory predispositions from brains to genomes

Memorizing and producing complex strings of sound are requirements for spoken human language. We share these behaviours with likely more than 4000 species of songbirds, making birds our primary model for studying the cognitive basis of vocal learning and, more generally, an important model for how memories are encoded in the brain. In songbirds, as in humans, the sounds that a juvenile learns later in life depend on auditory memories formed early in development. Experiments on a wide variety of songbird species suggest that the formation and lability of these auditory memories, in turn, depend on auditory predispositions that stimulate learning when a juvenile hears relevant, species-typical sounds. We review evidence that variation in key features of these auditory predispositions are determined by variation in genes underlying the development of the auditory system. We argue that increased investigation of the neuronal basis of auditory predispositions expressed early in life in combination with modern comparative genomic approaches may provide insights into the evolution of vocal learning.     

Hatching late in the season requires flexibility in the timing of song learning

Most songbirds learn their songs from adult tutors, who can be their father or other male conspecifics. However, the variables that control song learning in a natural social context are largely unknown. We investigated whether the time of hatching of male domesticated canaries has an impact on their song development and on the neuroendocrine parameters of the song control system. Average age difference between early- and late-hatched males was 50 days with a maximum of 90 days. Song activity of adult tutor males decreased significantly during the breeding season. While early-hatched males were exposed to tutor songs for on average the first 99 days, late-hatched peers heard adult song only during the first 48 days of life. Remarkably, although hatching late in the season negatively affected body condition, no differences between both groups of males were found in song characteristics either in autumn or in the following spring. Similarly, hatching date had no effect on song nucleus size and circulating testosterone levels. Our data suggest that late-hatched males must have undergone accelerated song development. Furthermore, the limited tutor song exposure did not affect adult song organization and song performance.