Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Mitochondrial rate variation among lineages of passerine birds

The order Passeriformes comprises the majority of extant avian species. Analyses of molecular data have provided important insights into the evolution of this diverse order. However, molecular estimates of the evolutionary and demographic timescales of passerine species have been hindered by a lack of reliable calibrations. This has led to a reliance on the application of standard substitution rates to mitochondrial DNA data, particularly rates estimated from analyses of the gene encoding cytochrome b (CYTB). To investigate patterns of rate variation across passerine lineages, we used a Bayesian phylogenetic approach to analyse the protein‐coding genes of 183 mitochondrial genomes. We found that the most commonly used mitochondrial marker, CYTB, has low variation in rates across passerine lineages. This lends support to its widespread use as a molecular clock in birds. However, we also found that the patterns of among‐lineage rate variation in CYTB are only weakly related to the evolutionary rate of the mitochondrial genome as a whole. Our analyses confirmed the presence of mutational saturation at third codon positions across the protein‐coding genes of the mitochondrial genome, reinforcing the view that these sites should be excluded in studies of deep passerine relationships. The results of our analyses have provided information that will be useful for molecular‐clock studies of passerine evolution.     

Song evolution,speciation, and vocal learning in passerine birds

Phenotypic divergence can promote reproductive isolation and speciation, suggesting a possible link between rates of phenotypic evolution and the tempo of speciation at multiple evolutionary scales. To date, most macroevolutionary studies of diversification have focused on morphological traits, whereas behavioral traits─including vocal signals─are rarely considered. Thus, although behavioral traits often mediate mate choice and gene flow, we have a limited understanding of how behavioral evolution contributes to diversification. Furthermore, the developmental mode by which behavioral traits are acquired may affect rates of behavioral evolution, although this hypothesis is seldom tested in a phylogenetic framework. Here, we examine evidence for rate shifts in vocal evolution and speciation across two major radiations of codistributed passerines: one oscine clade with learned songs (Thraupidae) and one suboscine clade with innate songs (Furnariidae). We find that evolutionary bursts in rates of speciation and song evolution are coincident in both thraupids and furnariids. Further, overall rates of vocal evolution are higher among taxa with learned rather than innate songs. Taken together, these findings suggest an association between macroevolutionary bursts in speciation and vocal evolution, and that the tempo of behavioral evolution can be influenced by variation in developmental modes among lineages.     

Evolution of sperm structure and energetics in passerine birds

Spermatozoa exhibit considerable interspecific variability in size and shape. Our understanding of the adaptive significance of this diversity, however, remains limited. Determining how variation in sperm structure translates into variation in sperm performance will contribute to our understanding of the evolutionary diversification of sperm form. Here, using data from passerine birds, we test the hypothesis that longer sperm swim faster because they have more available energy. We found that sperm with longer midpieces have higher levels of intracellular adenosine triphosphate (ATP), but that greater energy reserves do not translate into faster-swimming sperm. Additionally, we found that interspecific variation in sperm ATP concentration is not associated with the level of sperm competition faced by males. Finally, using Bayesian methods, we compared the evolutionary trajectories of sperm morphology and ATP content, and show that both traits have undergone directional evolutionary change. However, in contrast to recent suggestions in other taxa, we show that changes in ATP are unlikely to have preceded changes in morphology in passerine sperm. These results suggest that variable selective pressures are likely to have driven the evolution of sperm traits in different taxa, and highlight fundamental biological differences between taxa with internal and external fertilization, as well as those with and without sperm storage.     

Polyterritorialitytion in passerine birds

In some birds, males defend two spatially separated territories and attract females to each one in turn. it has been proposed that this behaviour, referred to as polyterritoriality, allows males to conceal their marital status, thereby deceiving females into accepting polygyny against their best interests. The deception hypothesis has gained general acceptance as an explanation for polyterritoriality in birds, but until recently the empirical basis for the hypothesis rested almost entirely on studies of Swedish populations of the pied flycatcher (Ficedula hypoleuca). New findings, emerging from further studies of the pied flycatcher and of other species, suggest several alternative selection pressures that may have favoured polyterritoriality.     

Latitude, elevation and body mass variation in Andean passerine birds

• 1 Relationships between body mass and latitude, and body mass and elevation are examined in the assemblage of Andean passerine birds.
• 2 Across species, body mass is positively correlated with the mid‐point of the species elevational distribution, but there is no significant relationship between body mass and latitudinal range mid‐point.
• 3 When the assemblage is separated into Andean endemic and non‐endemic species, the former group shows a significant positive relationship between body mass and elevation, and the latter a significant positive relationship between body mass and latitude (‘Bergmann’s rule’). Andean endemic species exhibit Bergmann’s rule once elevation is controlled for using multiple regression.
• 4 These relationships are not a consequence of the phylogenetic non‐independence of species. All the effects shown are very weak, with latitude and elevation explaining only a few per cent of the variation in body mass. Relationships are strongest when phylogenetically controlled analyses are performed just within genera.
• 5 The implications of these results for the mechanistic understanding of Bergmann’s rule are discussed.

     

Sperm length variation as a predictor of extrapair paternity in passerine birds

Background

The rate of extrapair paternity is a commonly used index for the risk of sperm competition in birds, but paternity data exist for only a few percent of the approximately 10400 extant species. As paternity analyses require extensive field sampling and costly lab work, species coverage in this field will probably not improve much in the foreseeable future. Recent findings from passerine birds, which constitute the largest avian order (∼5 900 species), suggest that sperm phenotypes carry a signature of sperm competition. Here we examine how well standardized measures of sperm length variation can predict the rate of extrapair paternity in passerine birds.

Methodology/Principal Findings

We collected sperm samples from 55 passerine species in Canada and Europe for which extrapair paternity rates were already available from either the same (n = 24) or a different (n = 31) study population. We measured the total length of individual spermatozoa and found that both the coefficient of between-male variation (CV_bm) and within-male variation (CV_wm) in sperm length were strong predictors of the rate of extrapair paternity, explaining as much as 65% and 58%, respectively, of the variation in extrapair paternity among species. However, only the CV_bm predictor was independent of phylogeny, which implies that it can readily be converted into a currency of extrapair paternity without the need for phylogenetic correction.

Conclusion/Significance

We propose the CV_bm index as an alternative measure to extrapair paternity for passerine birds. Given the ease of sperm extraction from male birds in breeding condition, and a modest number of sampled males required for a robust estimate, this new index holds a great potential for mapping the risk of sperm competition across a wide range of passerine birds.     

Phenotypic flexibility in passerine birds: Seasonal variation of aerobic enzyme activities in skeletal muscle

Improved winter cold tolerance is widespread among small passerines resident in cold climates and is generally associated with elevated summit metabolic rate (M_sum=maximum thermoregulatory metabolic rate) and improved shivering endurance with increased reliance on lipids as fuel. Elevated M_sum and improved cold tolerance may result from greater metabolic intensity, due to mass-specific increase in oxidative enzyme capacity, or increase in the masses of thermogenic tissues. To examine the mechanisms underlying winter increases in M_sum, we investigated seasonal changes in mass-specific and total activities of the key aerobic enzymes citrate synthase (CS) and β-hydroxyacyl CoA-dehydrogenase (HOAD) in pectoralis, supracoracoideus and mixed leg muscles of three resident passerine species, black-capped chickadee (Poecile atricapillus), house sparrow (Passer domesticus), and white-breasted nuthatch (Sitta carolinensis). Activities of CS were generally higher in winter than in summer muscles for chickadees and house sparrows, but not nuthatches. Mass-specific HOAD activity was significantly elevated in winter relative to summer in all muscles for chickadees, but did not vary significantly with season for sparrows or nuthatches, except for sparrow leg muscle. These results suggest that modulation of substrate flux and cellular aerobic capacity in muscle contribute to seasonal metabolic flexibility in some species and tissues, but such changes play varying roles among small passerines resident in cold climates.     

Cooperate or speciate: new theory for the distribution of passerine birds

In cooperatively breeding birds, adults often forego reproduction and help care for the offspring of others. A universal explanation for this mode of breeding has eluded evolutionary biologists, who have considered it to be a rare, and largely Australian, phenomenon. In a recent paper, Andrew Cockburn reports that the number of known cooperative breeders among oscine passerine birds has more than doubled since the last substantial review, published 16 years ago. Cooperative breeding is often the ancestral trait, and predominantly cooperative genera are species poor compared with their pair-breeding counterparts. Cockburn argues that speciation is less likely in cooperative clades, because the philopatric tendencies of individuals make them poor dispersers, colonizers and migrants. This new hypothesis helps explain the distribution and composition of migrant and island avifauna. However, a major challenge remains to reconcile the roles of phylogenetic history and current ecology in promoting cooperative behaviour.     

Body frontal area in passerine birds

Projected body frontal area is used when estimating the parasite drag of bird flight. We investigated the relationship between projected frontal area and body mass among passerine birds, and compared it with an equation based on waterfowl and raptors, which is used as default procedure in a widespread software package for flight performance calculations. The allometric equation based on waterfowl/raptors underestimates the frontal area compared to the passerine equation presented here. Consequently, revising the actual frontal areas of small birds will concomitantly change the values of the parasite drag coefficient. We suggest that the new equation     (m²) where m _B is body mass (kg) should be used when a value of frontal area is needed for passerines.     

Intraspecific variation in sperm length is negatively related to sperm competition in passerine birds

Spermatozoa are among the most diversified cells in the animal kingdom, but the underlying evolutionary forces affecting intraspecific variation in sperm morphology are poorly understood. It has been hypothesized that sperm competition is a potent selection pressure on sperm variation within species. Here, we examine intraspecific variation in total sperm length of 22 wild passerine bird species (21 genera, 11 families) in relation to the risk of sperm competition, as expressed by the frequency of extrapair paternity and relative testis size. We demonstrate, by using phylogenetic comparative methods, that between-male variation in sperm length within species is closely and negatively linked to the risk of sperm competition. This relationship was even stronger when only considering species in which data on sperm length and extrapair paternity originated from the same populations. Intramale variation in sperm length within species was also negatively, although nonsignificantly, related to sperm competition risk. Our findings suggest that postcopulatory sexual selection is a powerful evolutionary force reducing the intraspecific phenotypic variation in sperm-size traits, potentially driving the diversification of sperm morphology across populations and species.     

Comparative phylogeography and genetic structure of Vanuatu birds: control region variation in a rail, a dove, and a passerine

We examined variation in mitochondrial control region (Domain I) sequences in three distantly related species of birds found on multiple islands in Vanuatu: the Buff-banded Rail Gallirallus philippensis (n=21, 433bp), the Emerald Dove Chalcophaps indica (n=21, 513bp), and the Streaked Fantail Rhipidura spilodera (n=17, 326bp). Nucleotide and haplotype diversity were similar in all three species and showed little to no geographic structure within Vanuatu. Estimates of demographic parameters, tests for excess rare alleles, and the structure of both haplotype networks and pairwise mismatch distributions support a hypothesis of recent colonization and subsequent expansion in C. indica and R. spilodera but not in G. philippensis. Phylogenetic analyses suggest that G. philippensis samples from Vanuatu are polyphyletic, and further indicate that this species has had a more complex history of colonization than do the other two species. Estimated divergence times of alleles sampled in Vanuatu suggest there may be slight differences among species in the timing of colonization of Vanuatu despite differences in flight ability and presumed dispersal ability. In all three species, current populations probably derive from colonists that arrived several million years after the islands formed. Our comparisons illustrate the potential of direct genetic analyses of to highlight historical differences among co-distributed species with similar levels of phenotypic variation.     

Postexercise ketosis in night-migrating passerine birds

This study investigated the postexercise metabolism of six species of free-living, night-migrating passerine birds (European robin, pied flycatcher, wheatear, redstart, blackcap, and garden warbler). The birds were caught during autumn migration out of their nocturnal flight, and their metabolism changed from a fasting, highly active state to a fasting, resting state. Concentrations of six plasma metabolites of the fat, carbohydrate, and protein metabolism were measured during up to 10 h of recovery time. The metabolic changes indicated a biphasic pattern: (a) a quick first response to the reduced energy demands during the first 20 min of recovery, suggested by an increase and subsequent decrease of free fatty acid levels, and (b) subsequently, a postexercise ketosis and a reduction of lipolysis and proteolysis, suggested by high beta-hydroxy-butyrate and low free fatty acid, glycerol, triglyceride, and uric acid levels. This metabolic pattern differs from that of humans and rats, in which ketosis starts immediately postexercise or is absent in trained subjects. Since migrating birds are naturally adapted to endurance exercise, it is hypothesized that the high and long-lasting postexercise ketosis does not evoke physiological problems (such as hypoglycemia) but, by contrast, increases the ability of birds to rely on lipids, to a very high extent, during and after flight and decreases the dependence on glucose and glucogenic amino acids. Differences between species in fat stores and metabolic pattern support this hypothesis.     

Evaluating the contribution of dispersal to community structure in Neotropical passerine birds

For two centuries evolutionary biologists have sought to explain elevated biodiversity in the Neotropics. Although different process are known to be important, it is still not uncommon for researchers to emphasize a single mechanism. Recently, arguments have highlighted the importance of dispersal shaping community structure and evolution across the region. We examine this hypothesis by visualizing spatial variation in community structure for the majority of South American passerines (Aves) across the northern half of South America. By sampling over a contiguous area we show how community structure varies widely across Amazonia and surrounding regions. Our results support a combination of processes including: the inability of species to disperse across geographic barriers, Andean uplift, and variation in habitat type. Although dispersal is a factor, our results emphasize a lack of dispersal, driven primarily by features of the landscape, coupled with historical changes in climate to be important drivers of Neotropical diversity.     

Recent changes in body weight and wing length among some British passerine birds

We tested the prediction that global warming has caused recent decreases in body weight (Bergmann's rule) and increases in wing length (Allen's rule) in 14 species of passerine birds at two localities in England: Wicken Fen (1968–2003) and Treswell Wood (1973–2003).
Predicted long-term linear decreases in residual body weight occurred in four species: dunnocks (Wicken Fen), and great tits, blue tits and bullfinches (Treswell Wood). Non-linear decreases also occurred in reed warblers and blackcaps at Wicken Fen, which also had a surprising linear increase in residual body weight in blackbirds.
Residual wing lengths increased linearly, as predicted, in six of seven species at Wicken Fen. Whereas there were non-linear long-term increases in wrens, dunnocks and blackbirds in Treswell Wood. Unexpected linear decreases also occurred in residual wing lengths in willow warblers (Wicken Fen), and blue tits, great tits and chaffinches (Treswell Wood).
The most parsimonious explanation for such long-term changes in body weight is global warming, as predicted by Bergmann's rule. Greater site and species-specific effects on wing length (e.g. non-linear changes plus shorter wings in the woodland habitat) suggest a less straightforward conclusion concerning Allen's rule, probably because wing length involved variation in both bone and feather growth.
Changes in residual body weights and wing lengths often differed between species and were sometimes non-linear, perhaps reflecting short-term modifications in selection pressures. Human-induced influences are discussed, such as avian predator population densities and land-use change. Short-term variation in temperature had little effect, but rainfall did explain the unusual increase in blackbird body weight, possibly as a result of improving food (earthworm) availability.     

Seed depletion and landscape structure affect aggregative response in two wintering passerine birds

Capsule: Seed abundance but also seed depletion during winter modifies habitat use and aggregation behaviour of Eurasian Sky Lark Alauda arvensis and Meadow Pipit Anthus pratensis.

Aims: To investigate the effect of seed abundance on the density of two passerine birds, the Eurasian Sky Larks and the Meadow Pipit, at an agricultural landscape scale during winter.

Methods: Bird counts as well as seed density measurements were carried out in winter to quantify the aggregative response of farmland birds to seed abundance from November to March.

Results: Seed resources varied by a factor of ten between crop types, but declined so sharply over the winter that seed abundance was low and similar among crop types by late winter. Sky Larks selected for higher seed density plots, but only at the end of winter when the resources were the lowest. Conversely, Meadow Pipits did not show any aggregative response to seed abundance.

Conclusion: These results show that an uptake of seed-rich habitats in agricultural landscape would be very beneficial for wintering granivorous birds, by fulfilling the late winter ‘hungry gap’. The inclusion of seed mixtures in habitats that attract high densities of birds and retain seeds until late winter is of primary interest.     

DNA evolutionary rates in nine-primaried passerine birds

Differences in single-copy nuclear-DNA sequences among 13 species of passerine birds were measured using DNA-DNA hybridization. A matrix of pairwise dissimilarity values (delta mode distances) was constructed from analysis of fitted thermal dissociation curves. A least-squares method of phylogenetic estimation was used to construct two topologies from the distance matrix, one constraining branch lengths of sister taxa to be equal and the other permitting such lengths to vary. These topologies were identical in the pattern of branching of taxa, and the difference in their sums of squares was not statistically significant, suggesting that rates of DNA evolution in sister groups of nine- primaried oscines are equal. A nonparametric test for nonrandom variation in distances of sister groups to outgroup taxa revealed no statistically significant deviation from random variation that would be expected as a result of measurement error. However, the level of measurement error was such that rates of DNA evolution in sister taxa could vary by as much as 10% without being detected with the statistical methods used here.