Genetic divergences and intraspecific variation in corvids of the genus Corvus (Aves: Passeriformes: Corvidae) – a first survey based on museum specimens

The first comprehensive overview of intra‐ and interspecific variation within the genus Corvus as well as first insights into the phylogenetic relationships of its species is presented. DNA sequences of the mitochondrial control region were obtained from 34 of the 40 described species (including subspecies: 56 taxa). As the study was based mainly on museum material, several specimens did not yield the full length marker sequence. In these cases, only a short section of the control region could be analysed. Nevertheless, even these individuals could be assigned tentatively to clades established on the full length marker sequence. Inclusion of sequences of other corvid genera as available in GenBank clearly confirmed the monophyly of the genus Corvus. Within the Corvus clade several distinct subclades can be distinguished. Some represent lineages of single species or species pairs while other clades are composed of many species. In general, the composition of the clades reflects geographical contiguousness and confirms earlier assumptions of a Palearctic origin of the genus Corvus with several independent colonizations of the Nearctic and the Aethiopis. The Australasian radiation seems to be derived from a single lineage. The distribution of plumage colour in the phylogenetic tree indicates that the pale markings evolved several times independently. The white/grey plumage colour pattern – which is found also in other genera of the family Corvidae, for example, in Pica– occurs already in the species pair representing the first split within the genus Corvus (Corvus monedula, Corvus dauuricus). Thus, reversal to full black colour seems to have occurred as well. The use of colour traits as a phylogenetic marker within Corvus should be considered with severe caution.     

Isolation and characterization of microsatellite loci in a cooperatively breeding corvid,the American crow (Corvus brachyrhynchos)

We describe 11 microsatellite loci isolated from the American crow (Corvus brachyrhynchos), the most wide‐spread cooperatively breeding bird in North America. Polymorphism at these loci ranged from four to 43 alleles, and observed heterozygosities from 0.53 to 0.92. Genetic parentage analyses using these markers will allow us to describe the mating system of this common but socially complex species, and to interpret their behavioural interactions in light of relatedness.     

Analysing spatial and temporal variation in colony size: an approach using autoregressive mixed models and information theory

The spatial and temporal variation in population sizes of animal colonies are rarely studied simultaneously. I examined factors determining colony size (number of nests) for 23 colonies from the only breeding population of rook Corvus frugilegus in Spain over 7 years. Population sizes within colonies were highly predictable over time, with autocorrelations up to a distance (lag) of 6 years. Autoregressive mixed models were used to explain colony size as a function of environmental factors, while controlling for temporal autocorrelation. These factors included refuse tips, widely used as food resource, and a derived variable that incorporated the two factors most often related to avian colony size (inter-colony competition and foraging habitat around colonies). Autoregressive models provided a better fit to the data than models which did not consider temporal autocorrelation. The information-theoretic (AIC_c-based) approach revealed uncertainty in the selection of the best model explaining colony-size, but relatively strong support for certain variables. The highest weights of evidence were for year (ω _i  = 0.90) and the number of competitors per unit of foraging habitat (i.e., derived variable; ω _i  = 0.63), showing that the size of rook colonies in Spain was negatively affected by inter-colony competition relative to the foraging habitat surrounding the colonies. This variable measured within a 6-km radius from the colonies had ~30 times higher weight of evidence (more plausible) than the same variable measured within 3 km, indicating that food limitation may occur outside the breeding period. Sizes of colonies tended to decrease when distance between the colony and the nearest refuse tip increased. There was some evidence supporting the idea that the effect of the number of competitors per unit of foraging habitat on colony size varied from year to year, but statistical power was weak. These findings suggest that variation in number of rook nests within colonies reflects spatial and temporal heterogeneity of net food via both inter-colony competition and foraging habitat around the colony. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

北京麋鹿苑达乌里寒鸦迁徙和越冬期种群动态与集群行为的初步研究

2005年10月～2009年5月,在达乌里寒鸦(Corvus dauurica)迁徙和越冬期间,针对其在北京麇鹿苑内的种群数量变化,成体、幼体组成比例的变化,以及在麋鹿苑集群停歇的原因等问题,进行了调查分析.结果表明,达乌里寒鸦每年10月中旬前后开始到糜鹿苑聚集活动,次年5月份以后,集群现象消失.持续活动时间为(166±11)d,10月底或11月初种群数量达最大,2008年10月下旬,最大数量一度达(6 500±300)只,4个迁徙越冬期的最大种群数量存在极显著差异(X_((3))~ 2=8 484.47,P＜0.01＝.Krushkalwallis H检验表明,4个迁徙越冬季的种群数量之间差异不显著(X_((3))~2=6.14,P=0.11＞0.05).幼体的比例在秋季迁徙前期和春季迁徙后期相对较高,可达100%.观察表明,该鸟在麋鹿苑周边的垃圾场觅食后,选择在苑内饮水是其在麋鹿苑集群的主要原因.     

Duplicated growth hormone genes in a passerine bird, the jungle crow (Corvus macrorhynchos)

Molecular cloning, molecular phylogeny, gene structure and expression analyses of growth hormone (GH) were performed in a passerine bird, the jungle crow (Corvus macrorhynchos). Unexpectedly, duplicated GH cDNA and genes were identified and designated as GH1A and GH1B. In silico analyses identified the zebra finch orthologs. Both GH genes encode 217 amino acid residues and consist of five exons and four introns, spanning 5.2 kbp in GH1A and 4.2 kbp in GH1B. Predicted GH proteins of the jungle crow and zebra finch contain four conserved cysteine residues, suggesting duplicated GH genes are functional. Molecular phylogenetic analysis revealed that duplication of GH genes occur after divergence of the passerine lineage from the other avian orders as has been suggested from partial genomic DNA sequences of passerine GH genes. RT-PCR analyses confirmed expression of GH1A and GH1B in the pituitary gland. In addition, GH1A gene is expressed in all the tissues examined. However, expression of GH1B is confined to several brain areas and blood cells. These results indicate that the regulatory mechanisms of duplicated GH genes are different and that duplicated GH genes exert both endocrine and autocrine/paracrine functions.     

Habitat but not group size or recent predator activity affect corvid collective vigilance at carcasses

Vigilance is an important anti-predator behaviour that can be an indicator of the predation risk faced by potential prey animals. Here, we assess the collective vigilance, or the vigilance level of an entire group, of corvids (Family: Corvidae) at experimentally placed carcasses in a desert environment in Australia. Specifically, we explore the relationship between collective vigilance levels and the habitat in which the carcass was placed, the time since a potential predator (dingo Canis dingo, wedge-tailed eagle Aquila audax or red fox Vulpes vulpes) was present at a carcass, and the group size of corvids around the carcass. We found that corvids are more vigilant in open habitat, but that group size and the recent presence of a potential predator does not affect the collective vigilance behaviour of corvids. The results demonstrate the important link between habitat and vigilance, and that animals may adopt anti-predator behaviours irrespective of the size of the group in which they occur or the recent presence of a potential predator.     

Genetic signatures of intermediate divergence: population history of Old and New World Holarctic ravens (Corvus corax)

Many studies of phylogeography, speciation, and species limits restrict their focus to a narrow issue: gene tree monophyly. However, reciprocal monophyly does not provide an ideal touchstone criterion of any aspect of evolutionary divergence. There is a continuum of divergence stages as isolated populations go from initial allele frequency differences to well-differentiated species. Studying intermediate stages of divergence will increase our understanding of geographical speciation, species limits, and conservation priorities. We develop a conceptual framework and terminology for thinking about the stages of 'intermediate polyphyly'. The Holarctic clade of common ravens (Corvus corax), found throughout much of Eurasia and North America, provides a case study of these stages of intermediate divergence. We used coalescent, phylogenetic, and population genetic methods to investigate the history and current status of this Old World-New World distribution using 107 mitochondrial control region sequences. Phylogenetically, New World and Old World samples are intermixed. However, most samples are grouped into small subclades that are restricted to either the New World or the Old World, and only one haplotype is shared between the hemispheres. Analysis of moleculalr variance (amova) results reflect this low haplotype sharing between hemispheres (Phi(ST) = 0.13, P < 0.01). Isolation with Migration (im) coalescent results suggest a sustained period of divergence between the hemispheres and low levels of maternal gene flow. Although there has not been sufficient time to evolve reciprocal monophyly and some gene flow may occur, New World and Old World ravens are genetically quite distinct. We use this example to demonstrate these early stages of divergence as populations go from sharing only internal haplotypes, to sharing no haplotypes, to having population specific subclades. Studies of phylogeography, speciation and systematics will benefit from increased attention to these stages of intermediate polyphyly.     

Development of 10 microsatellite loci for Yellow-billed Magpies (Pica nuttalli) and corvid ecology and West Nile virus studies

We developed 10 polymorphic microsatellite loci for Yellow-billed Magpies (Pica nuttalli). The primers were tested across a population of 57 Central California Yellow-billed Magpies and displayed an average of 3.9 alleles per locus. Forty-one American Crows (Corvus brachyrhynchos) from California were polymorphic for seven of the loci with an average of 2.9 alleles per locus. One additional microsatellite-containing locus displayed diagnostic allele sizes and may be useful to distinguish between the two species. These corvid specific microsatellites will aid ecological studies of the population-level effects of diseases, such as West Nile virus.     

Dynamic species distribution models reveal spatiotemporal habitat shifts in native range‐expanding versus non‐native invasive birds in an urban area

Urbanisation as a major driver of changes leads to the extinction of some species while others increase in abundance, especially non‐native species. Spatiotemporal distribution patterns of these successful species are likely to be shaped by their response and tolerance to urban features. This study assesses the anthropo‐ecological requirements of two co‐occurring bird species, the native range‐shifting jackdaw Corvus monedula and the non‐native invasive ring‐necked parakeet Psittacula krameri. We built yearly models over an eight‐year period using an ensemble modelling approach assessing response differences through time and between species. Predictors describing human‐made structures, socio‐ecological proxies and resources availability were extracted from temporally coincident databases. Dispersal and habitat constraints were implemented in final models to provide more realistic forecasts of species future distributions. Ensemble models evaluated with a random partition of the training dataset showed a higher accuracy than those evaluated with an independent dataset from another time period. Our results highlight temporal variations in the relative importance of predictors for both studied species. Single‐season occurrence data may thus be insufficient to characterize species ecological requirements. The ring‐necked parakeet and the jackdaw showed different responses to urban features. Jackdaws preferred the more urbanized part of the city while the distribution of parakeets was strongly positively associated with the density of exotic ornamental trees. We concluded that ring‐necked parakeet range expansion is likely to be driven by its effective ability to exploit urban resources which native species do not or under exploit, suggesting an open window of foraging opportunities. However, the jackdaw may be misled by a high cavity availability and a large amount of low‐quality anthropogenic food in the urban core. We suggest that dynamic SDMs are a critical tool not only to forecast the future expansion of invasive species but also for a better understanding of processes driving urban biodiversity persistence.     

Problems of censusing Reed Buntings,Sedge Warblers and Reed Warblers

The daytime dispersal of Starlings from an urban winter communal roost was monitored by colour marking 1424 birds according to age and sex. Reports received from members of the public suggested that dispersal was homogeneous with respect to age and sex composition at all distances from the roost. There was evidence that dispersal was geographically heterogeneous with respect to sex composition, and this observation is discussed in relation to correlated variation in agricultural practices.