Genetic diversity of carrion and jungle crows from RAPD-PCR analysis data

RAPD-PCR analysis of the genetic diversity of the carrion crow (Corvus corone) and jungle crow (C. macrorhynchos) living in the continental parts of their species ranges and on some Russian and Japanese Far Eastern islands has been performed. Taxon-specific molecular markers have been found for each species. The genetic diversity of the carrion crow is considerably less than that of the jungle crow at the same genetic distance (P95 = 68.2%, DN = 0.27 and P95 = 88.4%, DN = 0.24, respectively). In both species, the genetic polymorphism of island samples is almost two times greater than that of continental samples (62 and 31.8%, respectively, for C. corone and 81.5 and 47.2%, respectively, for C. macrorhynchos). In addition, differences in genetic diversity between males and females (P95 = 55.1 and P95 = 72.1, respectively) has been found in the carrion crow but not in the jungle crow. The gene diversity of C. macrorhynchos is greater than that of C. corone: the mean numbers of alleles per locus are 2 and 1.81, effective numbers of alleles are 1.62 and 1.43, and the mean expected heterozygosities are 0.39 and 0.30, respectively. The phenograms and phylograms significantly segregate the clusters of the carrion and jungle crows. The clustering patterns of carrion crows corresponds to the intraspecies taxonomic and geographic differentiation: subspecies C. c. corone and C.c. orientalis living in the western and eastern parts of the species range, respectively, form different subclusters. The cluster of the jungle crow does not exhibit differentiation into subspecies C. m. mandshuricus and C. m. japonensis; molecular genetic differences between them are small.     

An analysis of population genetic differentiation and genotype–phenotype association across the hybrid zone of carrion and hooded crows using microsatellites and MC1R

The all black carrion crow ( Corvus corone corone ) and the grey and black hooded crow ( Corvus   corone cornix ) meet in a narrow hybrid zone across Europe. To evaluate the degree of genetic differentiation over the hybrid zone, we genotyped crows from the centre and edges of the zone, and from allopatric populations in northern (Scotland–Denmark–Sweden) and southern Europe (western–central northern Italy), at 18 microsatellites and at a plumage candidate gene, the MC1R gene. Allopatric and edge populations were significantly differentiated on microsatellites, and populations were isolated by distance over the hybrid zone in Italy. Single-locus analyses showed that one locus, CmeH9, differentiated populations on different sides of the zone at the same time as showing only weak separation of populations on the same side of the zone. Within the hybrid zone there was no differentiation of phenotypes at CmeH9 or at the set of microsatellites, no excess of heterozygotes among hybrids and low levels of linkage disequilibrium between markers. We did not detect any association between phenotypes and nucleotide variation at MC1R , and the two most common haplotypes occurred in very similar frequencies in carrion and hooded crows. That we found a similar degree of genetic differentiation between allopatric and edge populations irrespectively of their location in relation to the hybrid zone, no differentiation between phenotypes within the hybrid zone, and neither heterozygote excess nor consistent linkage disequilibrium in the hybrid zone, is striking considering that carrion and hooded crows are phenotypically distinct and sometimes recognised as separate species.     

Phylogenetic relationships of three species of crows (Corvidae, Aves) based on the restriction site variation of nuclear ribosomal RNA gene

Southern blot analysis of nuclear ribosomal DNA (rDNA) was carried out to examine phylogenetic relationships between three species of crows: Corvus cornix, C. corone and C. macrorhynchos. In this purpose DNA samples of birds were digested by 12 restriction enzymes (EcoRI, HindIII, PstI, BamHI, DraI, PvuII, KpnI, XbaI, BglII, BclI, SacI and AatI) and hybridized with the clones of mouse rDNA probes (18S, 28S and INT). Based on the data obtained and Gallus gallus restriction map as a standard the restriction site maps of the main rDNA repeating unit types (repetypes) were constructed. The length of crow rDNA genes was estimated to be 22.5 kb for Jungle and 22.0 kb for Hooded and Carrion crows. C. corone and C. cornix shared a common repetype which differed, by presence of two restriction sites (XbaI and PvuII) in the spacer region, from that of C. macrorhynchos with the estimated sequence divergence of 0.26%. Restriction-size variation was revealed between individuals of C. corone and C. cornix, although the substantial meanings of this variation remain unclear yet. These data suggest that the crow species evolve with slower rate of molecular evolution, as generally observed in other avian species, compared with the higher extent in external morphology, ecological features and behavior.     

Comparative phylogeography of two crow species: jungle crow Corvus macrorhynchos and carrion crow Corvus corone

The jungle crow Corvus macrorhynchos Wagler, 1827, and the carrion crow Corvus corone L., 1758, are two closely related species with similar ecological requirements that occupy wide distribution ranges in the Palearctic. We studied patterns of their genetic variation by using sequences of the mitochondrial cytochrome b gene. Corvus macrorhynchos demonstrates a low level of variation and differentiation throughout its range, except for a highly diverged population of Cheju Island (Korea). The haplotype network shows two haplogroups. The island group comprises populations of Sakhalin, Hokkaido, Honshu, and Kyushu, while the haplotypes of Taiwan and Ryukyu Islands proved to be closer to the mainland group, which also includes populations from the Primorye, Khabarovsk, Amur, and Magadan regions in the Russian Far East. This pattern allowed us to develop a phylogeographic hypothesis regarding the two modes of settling of the island populations. Concerning C. corone, the presence of two distinct haplogroups was confirmed within the range of C. c. orientalis. Both haplogroups are found within the same populations in Kamchatka and North Sakhalin, which implies secondary contacts there. Populations of C. corone are found to be rather stable in the western parts of its range, while in the Far East populations experienced recent growth, as was observed for C. macrorhynchos in general. The two species appear to have passed through different evolutionary scenarios.     

Ignorant hooded crows follow knowledgeable roost-mates to food: support for the information centre hypothesis

Communal roosting in birds may function to enhance foraging efficiency as explained by the information centre hypothesis, which predicts that successful foragers return from the roost to the rewarding food patch and that birds ignorant of this food follow knowledgeable roost-mates. We tested these predictions by exposing 34 radio-tagged, free-ranging, flock-living hooded crows (Corvus corone cornix) to a novel experimental set-up mimicking a superfluous food patch with maximum temporal and spatial unpredictability Each replicate lasted two days and was located on a new site. Data were collected during ten replicates over three years. First, a crow was more likely to visit the experimental food patch on the second day when it had been there on the first day. Second, when a crow had not been at this food patch on the first day, it was more likely to visit it on the second day if it had roosted together with a crow that had been there on the first day, but only if this knowledgeable roost-mate returned to the food patch on the second day. Our results support the information centre hypothesis and suggest that communal roosting might function to enhance foraging efficiency in hooded crows.     

History,environment and social behaviour: experimentally induced cooperative breeding in the carrion crow

Kin-based cooperative breeding, where grown offspring delay natal dispersal and help their parents to rear new young, has a long history in some avian lineages. Family formation and helping behaviour in extant populations may therefore simply represent the retention of ancestral features, tolerated under current conditions, rather than a current adaptive process driven by environmental factors. Separating these two possibilities challenges evolutionary biologists because of the tight coupling that normally exists between phylogeny and the environmental distribution of species and populations. The carrion crow Corvus corone corone, which exhibits extreme interpopulational variation in the extent of cooperative breeding, with populations showing no delayed dispersal and helping at all, provides a unique opportunity for an experimental approach. Here we show that offspring of non-cooperative carrion crows from Switzerland will remain on the natal territory and express helping behaviour when raised in a cooperative population in Spain. When we transferred carrion crow eggs from Switzerland to Spain, five out of six transplanted juveniles delayed dispersal, and two of those became helpers in the following breeding season. Our results provide compelling experimental evidence of the causal relationship between current environmental conditions and expression of cooperative behaviour.     

Morphology of the Hooded crow Corvus corone cornix in relation to age, sex and latitude

The morphological variation of the Hooded crow at Trondheim, Norway, was studied, based on a sample of 734 birds collected during a six year period. Mouth colour, plumage colour, skull thickness and feather length were found to be characters which could readily be used to separate juveniles from adults. Females aged 15–19 months had a thinner skull roof than older female birds. Low coefficients of variation were found for the lengths of the third primary and of the tail feathers. A discriminant analysis showed that of the various body dimensions studied bill height and bill length distinguished the sexes most precisely. A high degree of sexual dimorphism was also found to exist in body weight and in the thickness of the skull roof.
Those body structures which develop at about the same stage during the juvenile growth period were associated with the same principal component, viz. the lengths of bony structures, parts that develop early on in life, were intercorrelated (wing bones, tarsus, bill basis and the width of foramen magnum). The lengths of the primaries and of tail feathers were also intercorrelated, structures which develop late.
The mean body weight of the Hooded crow population studied in Norway was intermediate between that of the Hooded crow in Germany and of the Carrion and Hooded crows in England and Scotland. No such differences were found in wing length. Norwegian Hooded crows have shorter tails than German ones, but their bills are much larger, in particular for the females. Therefore, the degree of sexual dimorphism in bill size seems to be reduced at high latitudes.     

Tolerance and Social Facilitation in the Foraging Behaviour of Free‐Ranging Crows (Corvus corone corone; C. c. cornix)

Social foraging provides animals with opportunities to gain knowledge about available food. Studies indicate that animals are influenced by social context during exploration and are able to learn socially. Carrion and hooded crows, which are opportunistic generalists with flexible social systems, have so far received little focus in this area. We combined observational and experimental approaches to investigate social interactions during foraging and social influences on crow behaviour within a free‐ranging population at Vienna Zoo, which included 115 individually marked crows. We expected the crows to be tolerant of conspecifics during foraging due to high food abundance. We predicted that social context would enhance familiar object exploration, as well as a specific foraging strategy: predation by crows on other species. We found that crows were highly tolerant of one another, as reflected by their high rates of cofeeding – where they fed directly beside conspecific(s) – relative to affiliative or agonistic interactions. Evidence for social facilitation – when the observer's behaviour is affected by the mere presence of a model – was found in both object exploration and predation behaviour. Specifically, crows touched the objects more frequently when others were present (whilst only approaching the objects when alone), and conspecifics were present more frequently during predation events involving the high‐risk target species. Evidence for enhancement during object exploration – where the observer's attention is drawn to a place or object by a model's actions – was not confirmed in this context. Our results highlight the role played by the presence of conspecifics across different contexts: natural foraging behaviour, familiar object exploration and a specific foraging strategy. To our knowledge, this is one of the first corvid studies aimed at teasing apart specific social influence and learning mechanisms in the field. These crows therefore make promising candidates for studying social learning and its consequences under naturalistic conditions.     

On the phylogenetic relationship of Corvinae birds (Aves, Corvidae) from data of partial sequencing of cytochrome b gene mitochondrial DNA

To establish phylogenetic relationships within the corvine birds at the interspecific and intergeneric levels, the sequence of the mitochondrial DNA cytochrome b gene was analyzed. The NJ, UPGMA, and MP trees showed similar clustering. Relationships between the jungle crow, on the one hand, and the rook and Australian raven, on the other hand, were closer than between the jungle crow and the hooded and carrion crows. Mitochondrial genome of Australian raven displayed the closest similarity to the ancestral genome of the genus Corvus. Populations inhabiting the eastern part of the carrion crow C. corone orientations area were statistically significantly subdivided into three lineages. These data also confirmed the hypothesis on the location of the carrion crow ancestral lineage in the southeastern part of the area. In general, the transition and transversion substitution levels, their relationships, and distribution over codon positions were similar to that already reported for birds. Synonymous transitions in the third codon position were the prevailing substitution type. Using standard calibration scales, the time of divergence between species and genera within the corvine family was estimated to be 3.1-4 and 3.8-8.8 Myr, respectively. The divergence time between the examined corvine birds and birds of paradise constituted from 8 to 10 Myr.     

Phylogeographic carrion, hooded and jungle crows (Aves, Corvidae) from data on partial sequencing of the mitochondrial DNA cytochrome B gene

Distribution of mitochondrial DNA cytochrome b gene haplotypes in two crow species was examined by means of sequencing of the 336-bp gene fragment. The topology of the NJ and UPGMA trees showed that the carrion crow range was split into two parts due to the presence of significantly diverged ancestral lineage localized in the southeastern part of the range. The carrion crow populations, inhabiting a territory ranging from France to northern Sakhalin, along with interspersed hooded crow populations and hybrid Siberian populations, shared a common haplotype. The border between two carrion crow lineages revealed is located in central Sakhalin. The subdivision of two weakly differentiated lineages within the jungle crown range, also observed within this territory, coincided with the subspecies division of this species. The estimated genetic distances indicate the isolation of the subgenus Coloeus. These data also suggest the convergent similarity between the chough Pyrrhocorax pyrrhocorax and the Corvus genus, as well as the conspecificity of Corvus corone corone and C. c. cornix.     

Genetic variation of carrion and hooded crows and their hybrids based on RAPD-PCR data

RAPD-PCR analysis of genetic variability of carrion crows, hooded crows and their phenotypic hybrids from the zones of overlap of their areals and hybridization of parental forms in Siberia has been carried out. According to RAPD variability the following parameters were estimated for both species and their hybrids: genetical distances (DN), polymorphism (P95), mean expected heterozygosity (H(e)), gene fixation coefficient (Fst) and the portion of interpopulation differentiation (Gst). It was shown that the rate of genetic variability in carrion crows and hooded crows is less than in the hybrids. It turned out that the hybrids were genetically closer to each other (DN = 0.295) than to both parental species (DN = 0.441 and 0.397 with C. corone and C. cornix, correspondingly). The studied individuals of C. cornix show the minimal level of heterozygosity and polymorphism (H(e) = 0.12 and P95 = = 27.8%) in comparison with C. corone and the crows of hybrid zone (H(e) = 0.18 and 0.20; P95 = 42.6% and 50.4%, correspondingly). Right's indexes of inbreeding (Fst = 0.285) and population spliting (Gst = 0.352) indicate low genetic variability in all the analysed birds. Nevertheless UPGMA and NJ dendrogrammes differentiate the individuals from the areals of carrion crows, hooded crows and from hybrid zone to different clusters. Our data correlate with molecular-genetic investigations of Corvidae carried out with other methods, and do not contradict the conception of carrion crows and hooded crows as semispecies in the context of the conception of overspecies.     

Behavioural Type Affects Space Use in a Wild Population of Crows (Corvus corone)

While personality‐dependent dispersal is well studied, local space use has received surprisingly little attention in this context, despite the multiple consequences on survival and fitness. Regarding the coping style of individuals, recent studies on personality‐dependent space use within a habitat indicate that ‘proactive’ individuals are wider ranging than ‘reactive’ ones. However, such studies are still scarce and cover limited taxonomic diversity, and thus, more research is needed to explore whether this pattern generalises across species. We examined the link between coping style and space use in a population of crows (Corvus corone) freely inhabiting the urban zoo of Vienna, Austria. We used a binary docility rating (struggle during handling vs. no struggle) and a tonic immobility test to quantify individual coping style. Individual space use was quantified as the number of different sites at which each crow was observed, and we controlled for different number of sightings per individual by creating a space use index. Only the binary docility rating showed repeatability over time, and significantly predicted space use. In contrast to previous studies, we found that reactive crows (no struggle during handling) showed wider ranging space use within the study site than proactive individuals (who struggled during handling). The discrepancy from previous results suggests that the relationship between behavioural type and space use may vary between species, potentially reflecting differences in socioecology.     

Risk assessment by crow phenotypes in a hybrid zone

Predation is one of the most selective forces in evolution and, thus, predation may select against hybrids in narrow hybrid zones. It may be possible that parental phenotypes and hybrids differ in their responses towards predators or humans. As predation is difficult to observe I used flight-initiation distance (FID) as a metric of risk assessment. FID is a measurable outcome of the trade-off between fleeing and remaining. Here, I tested whether hybrid and parent crow phenotypes (Corvus corone, Corvus cornix) from the hybrid zone in Eastern Germany differ in their FID. Further, I measured many environmental and social variables to control statistically for their influence on FID. I sampled 154 individuals (53 hooded crows, 54 carrion crows, and 48 hybrids) in the hybrid zone in eastern Germany. I calculated a general linear model using a stepwise backward procedure to establish a minimum model containing only significant variables that explained FID in crows. The variable phenotype (hooded, carrion, hybrid) was then added to the model. There were no differences in FID between hybrids and both parental phenotypes types, suggesting similar risk assessment. This suggests that hybrids may behave similarly in their decision to flee as their parent phenotypes, which, in turn, provides no evidence for a selective disadvantage. An additional analysis focusing on pure phenotypic flocks showed that hybrids in pure hybrid flocks had a lower FID than both parental species in pure flocks. This suggests that hybrids in pure hybrid flocks may be at a disadvantage.     

Direct fitness benefits of group living in a complex cooperative society of carrion crows, Corvus corone corone

The social behaviour of carrion crows varies between populations. In northern Spain cooperatively breeding groups form through delayed natal dispersal and/or immigration of individuals (usually males) into the territory. In this population, carrion crows therefore breed as either unassisted pairs, pairs with nondispersing 1-2-year-old helpers (nondispersers), pairs with immigrant helpers or mixed groups (pairs with both immigrants and nondispersers). We used a microsatellite-based genotyping system to determine the parentage of 57 nestlings (19 broods). Polygamous mating was involved in 26% of the broods and reproduction was shared among group members of both sexes in at least three groups. Immigrants of both sexes can therefore gain access to mates by living in a group, while reproduction is unlikely to involve nondispersers. This implies that nondispersers and immigrants gain different sorts of benefits from group living and helping at the nest. Our genetic data confirmed that nondispersers associated with their parents on the natal territory and therefore that delayed natal dispersal leads to family formation in the carrion crow. Polygamous mating was not found in groups without immigrants, suggesting that, in this population, breeders lose parentage in their brood when sociality is extended beyond the limit of the nuclear family.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Growth and morphology of nestling Hooded crows Corvus corone cornix, a sexually dimorphic bird species

The growth and morphology of nestling Hooded crows, Corvus corone cornix , were studied at Trondheim, Central Norway, during 1980 and 1981. The sexual size dimorphism of the nestlings developed while they were in the nest, but was less pronounced than that of the juvenile birds. Female young grew relatively faster than the males. The size variation of the morphological characters studied was greater in the males than in the females, during the entire nestling period. The order of development of the different morphological characters seemed to be adapted to the needs of the nestlings during the nestling period. Growth of the bone and bill characters was the highest priority, body weight and feather characters the lowest. In general, the sizes of the different morphological characters were highly intercorrelated, particularly those that developed at the same time during the growth period. In the favourable breeding season of 1981, the nestlings were larger than in the unfavourable season of 1980. The high priority characters of the males increased more in size than did those of the females, and the low priority characters of the females increased more in size than did those of the males. About 75% of the nestlings could be correctly sexed from their external morphological characteristics. The findings support the view that the parental cost of rearing a male offspring is greater than that of a female.     

Relative numerousness judgements in crows and pigeons in the urgent comparison of stimuli earlier linked to different amounts of reinforcement]

The ability of pigeons (Colomba livia, L.) and crows (Corvus corone cornix, L.) was studied to realize urgent numerousness judgement of reinforcement consisting of discrete elements (wheat grains and meal worm larvae, respectively). In the process of preliminary training the birds mastered the information about the conformity of the feeder colour with the definite number (1-9 for pigeons and 5-12 for crows) of reinforcement units at isolated presentation of feeders. In test at presentation of pairs formed from these feeders, pigeons and crows chose the stimulus connected with a greater quantity of reinforcement. In the range of 1-8 units the precision of choice in pigeons depended on absolute and relative differences between comparing values. In crows in the range of 6-12 this dependence was not revealed. The ability to solve the given test is considered as one of manifestations of elementary reasoning.     

Why do zoos attract crows? A comparative study from Europe and Asia

Crows have successfully colonized many cities, and urban zoos have been important in this process. To evaluate why zoos attract crows, we quantified crow numbers and behavior in three zoos in Europe (Debrecen, Edinburgh, Vienna) and one in Asia (Sapporo). Data were collected in 445 surveys over 297 days in summer 2014 and winter 2014–2015. We found that crow numbers were highest in Vienna, intermediate in Debrecen and Edinburgh and lowest in Sapporo, increased significantly from summer to winter (Debrecen, Edinburgh, Vienna), and from mornings to afternoons (Debrecen, Sapporo, Vienna), and were higher in sunny weather than in cloudy weather with precipitation and when visitor numbers were low (Debrecen, Vienna). The crows' use of natural food was highest in Vienna, intermediate in Edinburgh and Sapporo, and low in Debrecen. The use of anthropogenic food was high in Debrecen and Sapporo, where the availability of open grassy areas typically used by crows for natural foraging was low. In Sapporo, food availability was more limited than in other zoos, resulting in strong territoriality and few crows in summer, which decreased further in winter. Our study indicates that crows are primarily attracted to zoos by food availability and secondarily by breeding opportunities and that the relative importance of natural versus anthropogenic food sources may vary with zoo habitat structure. Our study draws attention to a previously overlooked role of zoos in urban biodiversity conservation. It may also provide useful information for the management of crow populations, if necessary, and for the planning of urban areas.     

Hybrid zone maintenance by non-adaptive mate choice

In a spatial simulation we show that formation of a stable, narrow hybrid zone requires no selection, only that individuals can recognize their own population type since this is a prerequisite for assortative mating. The European crow occurs in two species/subspecies that meet in a long hybrid zone, the black carrion crow Corvus corone and the grey and black hooded crow C. cornix. In a previous study (Brodin A, Haas F (2006) Anim Behav 72:139) we mimicked sexual imprinting in nestling crows with artificial neural networks, using a learning process that simulates proximate retina perception. The networks were trained on 700 images to recognize either carrion, hooded or hybrid phenotypes as their own. After training the networks were exposed to 300 new, unfamiliar pictures of crows. The networks that had been trained on pure subspecies then showed strong preference for their own type. Networks trained on hybrid crows showed weaker preference for their own type. Assuming that these preferences will determine mate choice preferences we have combined them with empirical data on dispersal of young crows to investigate what kind of geographical distribution pattern this would create. We then assume that the two subspecies meet and hybridize along a straight border. In only 60 generations a stable, narrow hybrid zone that resembles the real hybrid zone was formed. This zone remained stable over time. With minor adjustments of parameters the results would not only fit the width but also the shapes of the clines in the field. An erratum to this article can be found at     

Attitudes toward and Acceptability of Management Strategies for a Population of Hooded Crows (Corvus cornix) in Slovenia

Recently, the number of hooded crows (Corvus cornix) in Slovenia has increased, which has resulted in a higher number of situations perceived as human–wildlife conflicts. The purpose of our survey was to provide a basis for population management measures of hooded crows that would be both feasible and acceptable to the public. An online survey based on snowball sampling received 1,042 responses, representing about 0.05% of the total and 0.13% of the active human population of the Republic of Slovenia. The major findings are that the majority of respondents were interested in learning more about problems caused by crows and think that crow numbers should be reduced, but by measures that would not cause their death. About one-third of respondents would be willing to participate in measures directed to controlling the crow population; however, about one-third would actively oppose such measures. They would even sign a petition regarding control measures. From the results, we conclude that the most suitable strategies in terms of acceptability would be passive, by educating people about how to avoid conflict with crows, and by better management of the resources affecting their foraging success.     

Mycoplasmosis in captive crows and robins from Minnesota

Mycoplasma sturni is a recently described organism previously associated with conjunctivitis in European starlings (Sturnus vulgaris), northern mockingbirds (Mimus polyglottos) and blue jays (Cyanocitta cristata). Herein we describe the isolation of M. sturni from an American crow (Corvus brachyrhynchos) presenting with conjunctivitis. A nested-PCR was designed for identification of M. sturni in clinical specimens and the sensitivity of the reaction was found to be 10 colony-changing units. The organism was found in asymptomatic American crows caged with a nestmate of the crow with conjunctivitis. Mycoplasma sturni also was found in asymptomatic American robins (Turdus migratorius) and in a European starling (Sturnus vulgaris) housed at the same facility as the crows. Heterogenity of M. sturni isolates from different host species was found by random amplified polymorphic DNA (RAPD) analyses. Heterogeneity also was found among M. sturni isolates recovered from American crows. We suggest that M. sturni can successfully infect American crows and American robins with or without the presence of clinical disease. Furthermore, we demonstrate that nested-PCR is an effective method for the detection of M. sturni and that substantial genetic heterogeneity exists among natural isolates of this bacterial pathogen.