The Crow Corvus corone hybrid zone in southern Denmark and northern Germany

In hybrid zones genetically differentiated populations meet and interbreed. As they result from ongoing divergence and potential speciation, such zones provide opportunities to study how different factors affect this process. One well-known avian hybrid zone is that between the Hooded Crow Corvus corone cornix and Carrion Crow C. c. corone , which extends throughout much of Europe. We compare the current position and width of the zone in southern Denmark and northern Germany with that described in detail by Meise in 1928. A GPS technique was used to determine the geographical position of more than 1000 Crows in established pairs in which all individuals were classified as either Carrion, Hooded or hybrid phenotype. The data show that the Carrion Crow has expanded northwards and that this is most pronounced at the eastern side of the study area. Here the hybrid zone has moved around 19 km. However, the width of the zone seems to have remained stable over time. Possible reasons for why the zone may have moved are discussed.     

Selektive Partnerwahl der Aaskr?he (Corvus corone) in der Hybridisierungszone von Rabenkr?he (C. c. corone) und Nebelkr?he (C. c. cornix)

Zusammenfassung Die als Unterarten klassifizierten europäischen Formen der Aaskrähe, Rabenkrähe und Nebelkrähe, besiedeln verschiedene, aneinandergrenzende Verbreitungsgebiete und hybridisieren in der Kontaktzone. Die Nachkommen von Mischpaaren sind fruchtbar und können sowohl mit anderen Hybriden als auch mit Raben- und Nebelkrähen erfolgreich brüten. Trotzdem kommt es zu keiner völligen Vermischung der Formen und/oder Verlagerung der Verbreitungsgebiete. Vor diesem Hintergrund untersuchten wir die Partnerwahl von Aaskrähen in der Hybridisierungszone auf der nordfriesischen Insel Amrum und stellten fest, daß Partner gleichen Phänotyps häufiger miteinander verpaart waren, als stochastisch zu erwarten gewesen wäre. Unsere Daten bestätigen vergleichbare Studien aus Hybridisierungszonen in Italien und Sibirien. Wir schließen daraus, daß phänotypisch selektive Partnerwahl bei der Aaskrähe ein allgemeines Phänomen sein könnte und diskutieren, warum dieses Verhalten anfitness-relevante Parameter gekoppelt sein sollte. Um welche es sich dabei handeln könnte, wurde bisher nicht hinreichend untersucht und muß deshalb offen bleiben.

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Non-random mating of Crows (Corvus corone) in a hybrid zone of Carrion Crow (C. c. corone) and Hooded Crow (C. c. cornix)

Summary Carrion Crow and Hooded Crow are regarded as subspecies of the Crow. They show frequent hybridisation along the adjacent borders of their distribution. Mixed pairs produce fertile offspring which are able to breed successfully with both hybrids and mates of either phenotype. However, hybridisation does not lead to phenotypic changes of Carrion and Hooded Crows in general nor in their distinct distribution. We studied the mating behaviour of Crows in the hybrid zone on the Island of Amrum (Schleswig-Holstein, Germany) and found evidence that Crows may prefer mates of the same phenotype. Our data confirm previous studies which reported assortative mating with respect to plumage coloration from hybrid zones in Italy and Siberia. We discuss why this behaviour should be related tofitness traits which in our opinion have not yet been studied adequately nor identified.

     

Mussel dropping by Carrion and Hooded crows: biomechanical and energetic considerations

Dropping live mussels (Mytilus sp.) onto hard substrata by Carrion Crows (Corvus corone) and Hooded Crows (Corvus cornix) to access their flesh is a commonly observed behavior from late summer to spring in the United Kingdom and Ireland. Despite previous studies, several aspects of prey‐dropping behavior remained incompletely understood. From September 2008 to January 2010, we determined the heights of drops, likelihood of shell breakage from drops at different heights, effect of mussel size on breakability, energetic costs of flying to drop heights, and the energetic costs of transporting mussels from mussel beds to dropping sites. We studied Carrion Crows on the Isle of Cumbrae, Scotland, and Hooded Crows in Cork Harbor, Ireland. Initial experiments were carried out with mussels to determine breakability in relation to size and drop height, and to estimate mussel energy content. Sizes of mussel shells at Hooded Crow dropping sites were compared with those of live mussels from source mussel beds. Adult Carrion Crows (N = 10) dropped mussels from a mean height of 4.7 m, and adult Hooded Crows (N = 21) from 4.8 m. These heights were close to the minimum (4–4.8 m; determined experimentally) required to break all mussel shells on the first drop. Dropping mussels from the minimum height that guarantees breakage reduces handling time and, by minimizing the size of the resulting debris field, likely reduces the risk of kleptoparasitism. Juvenile Hooded Crows (N = 13) dropped mussels onto suboptimal substrates (gravelly mud) from variable heights (mean = 6.1 m) with a low success rate (0% on first drop). This inefficiency could reflect either inexperience or exclusion from prime hard‐substrate dropping sites by adults. Foraging Hooded Crows selected larger mussels, dropping no mussels <32‐mm shell length. Energetic calculations indicate that a Hooded Crow lifting a medium‐sized mussel (55‐mm shell length) to a height of 5 m incurs a cost of only 0.3% of energy assimilated from that mussel, whereas travel to and from a mussel bed 200 m away costs 5.8% of that energy. These results suggest that choice of mussel dropping height by crows is determined by shell breakability rather than the cost of flying up to the dropping height.     

Aprocta matronensis in crows (Corvus corone corone) from northern Italy.

Aprocta matronensis (Nematoda, Spirurida) has been found in the orbital cavities of carrion crows. Hooded crows and crows from the hybrid crow zone in the studied areas were not infected. The parasite population showed the typical morphology described for this species. This is the first record of A. matronensis in crows in Italy and in Corvus corone corone. Some hypotheses about the distribution of the parasites in crow populations are discussed.     

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.     

Heterochromatin heterogeneity and rapid divergence of the sex chromosomes in Chorthippus parallelus parallelus and C. p. erythropus (Orthoptera).

Mitotic chromosomes from individuals of a Pyrenean hybrid zone between two subspecies of the grasshopper Chorthippus parallelus have been pretreated as for C-banding and subsequently stained with 4,6-diamidino-2-phenylindole (DAPI) and (or) Chromomycin A3 (CMA). The results, which are different from those obtained with Giemsa C-banding or standard DAPI - CMA treatments show (i) hidden heterochromatic heterogeneity that may be correlated with the existence of distinct families of repetitive DNAs, (ii) information about the possible independent origin of the three detected types of heterochromatin, and (iii) a further marker difference between the sex chromosomes of these two subspecies. This last result leads us to discuss the possible differential rates of evolution of sex chromosomes and autosomes in these subspecies and provides us with a new tool for the study of the structure and dynamics of this hybrid zone.     

Nest site selection and patterns of nest re-use in the Hooded Crow Corvus cornix

Capsule: Hooded Crows Corvus cornix selected nesting trees based on species, height, grouping and distance from an occupied house. Nest re-use was common and pairs that re-used old nests produced more fledglings than those that built a new nest.

Aims: To determine the features of trees that influenced whether they were used by Hooded Crows as nest sites, to establish what factors influenced nest re-use between years and to explore potential costs or benefits of nest re-use.

Methods: In a large area of Orkney, Scotland, the features of trees that contained a Hooded Crow nest were compared to those of trees where nests were absent. Patterns of nest re-use between years were examined in relation to the availability of alternative sites, previous nesting success and the number of equivalent options to the tree used previously within 200?m of this site.

Results: Hooded Crows favoured spruce and pine trees as nest sites, above the most locally abundant tree species, elder and willow. Preference for trees increased with tree height, local tree density and distance from occupied houses. Over half of the crows studied re-used an old nest when one was available and crows that re-used an old nest fledged more offspring than those that built a new nest. The likelihood of a new nest being built increased as the number of potential locations to build increased. Territories where a nest survived the winter were more likely to be reoccupied the following year than those where nests fell, while territories with fewer trees around the old site were most likely to be abandoned, suggesting that those were territories of lower quality.

Conclusions: Hooded Crows displayed strong preferences for nest sites that might favour nesting success by offering concealment, shelter and protection from ground-based predators. Nest re-use was common, especially when alternative sites were scarce, and appeared to facilitate greater reproductive output.     

Chromosomal differentiation through an Alpine hybrid zone in the grasshopper Chorthippus parallelus

The grasshopper Chorthippus parallelus is genetically differentiated over its range into at least five major geographical subdivisions. Two of these subdivisions, designated as subspecies, meet and form a hybrid zone in the Pyrenees. These subspecies differ for a wide range of morphological, behavioural and chromosomal characters, which vary clinally across the zone. A further geographical subdivision exists within Italy. Here we present the first evidence for chromosomal divergence between populations of the grasshopper north and south of the Alps. Across two Alpine cols (Col de Larche, between France and Italy; Passo de Resia, between Austria and Italy), these populations differ in both structural and functional components of the X-chromosome. Northern Alpine individuals possess an active nucleolar organizing region (NOR) at the distal end of the X-chromosome and an associated region of heterochromatin (C-band). Both these features are absent from individuals from the south of the Alps. However, all individuals examined carry distally located rDNA on the X-chromosome. Clinal transition was examined in the distal C-band in transects through the two cols. The cline centres are roughly coincident with the tops of the cols. Both clines were of similar widths (Col de Larche, 21.88 km; Passo de Resia, 24.05 km), and therefore much wider than those for an X-linked distal C-band in the Pyrenean hybrid zone. This suggests that there are different selective pressures on the cytogenetic characters in the Alps. The results are discussed in the context of the historical population dynamics of the species in relation to the climatic changes associated with the Pleistocene ice ages.     

New evidence for hybrid zones of forest and savanna elephants in Central and West Africa

The African elephant consists of forest and savanna subspecies. Both subspecies are highly endangered due to severe poaching and habitat loss, and knowledge of their population structure is vital to their conservation. Previous studies have demonstrated marked genetic and morphological differences between forest and savanna elephants, and despite extensive sampling, genetic evidence of hybridization between them has been restricted largely to a few hybrids in the Garamba region of northeastern Democratic Republic of Congo (DRC). Here, we present new genetic data on hybridization from previously unsampled areas of Africa. Novel statistical methods applied to these data identify 46 hybrid samples – many more than have been previously identified – only two of which are from the Garamba region. The remaining 44 are from three other geographically distinct locations: a major hybrid zone along the border of the DRC and Uganda, a second potential hybrid zone in Central African Republic and a smaller fraction of hybrids in the Pendjari–Arli complex of West Africa. Most of the hybrids show evidence of interbreeding over more than one generation, demonstrating that hybrids are fertile. Mitochondrial and Y chromosome data demonstrate that the hybridization is bidirectional, involving males and females from both subspecies. We hypothesize that the hybrid zones may have been facilitated by poaching and habitat modification. The localized geography and rarity of hybrid zones, their possible facilitation from human pressures, and the high divergence and genetic distinctness of forest and savanna elephants throughout their ranges, are consistent with calls for separate species classification.     

Vocal mimicry of hosts by Great Spotted Cuckoo Clamator glandarius: further evidence

Great Spotted Cuckoo nestlings were shown, after some days in the nest, to have begging calls that differed depending on whether they were being reared by Magpies or Carrion Crows. They also produced calls of a pitch and repetition rate that implied a high level of hunger.     

Strong premating divergence in a unimodal hybrid zone between two subspecies of the house mouse

Although selection against hybridization is expected to generate prezygotic divergence in unimodal hybrid zones, such a pattern has been seldom described. This study aims to better understand how prezygotic mechanisms may evolve in such zones. We investigated prezygotic divergence between populations of two subspecies of mice (Mus musculus musculus and M. m. domesticus) located at the edges of their unimodal hybrid zone in Denmark, and we developed an original multiple-population choice-test design, which allows assessment of within and between subspecies variation. Our study demonstrates that a strong assortative preference characterises one of the two subspecies (musculus) and that urinary signals are involved in this subspecies recognition. Taking into account the specific genetic and geographical characteristics of the Danish hybrid zone, we discuss the influence of the above pattern on its fate and the mechanisms that could have favoured this prezygotic divergence, among which the role of recombined populations constituting the core of the zone.     

Birding and DNA: species for the new millennium

Capsule Based on the 1999 Witherby Memorial Lecture – reviews how developments in molecular and population genetics have led to a reappraisal of species limits in birds.

The taxonomy of birds of the West Palearctic has moved from the comparative stability of the ‘Voous List’ into a period of serious activity, with new data emerging in almost every issue of every evolutionary and avian journal! This activity comes from two directions. Firstly, developments in population genetics, molecular biology, acoustics, behaviour and distributional studies have opened new avenues to measuring differentiation among groups of birds. This, in turn, has led to the recognition that earlier views of what constitutes a ‘species’ are in need of modification (‘improvement’), and the emergence of the ‘lineage concept’ of species. I review some of the species concepts most relevant to avian studies, and attempt to show how and why this change has happened, and its consequences for taxonomy and species limits. Examples are given in the form of ‘case studies’, and include Carrion/Hooded Crows Corvus corone/cornix, Green-winged/Eurasian Teals Anas carolinensis/crecca and Phylloscopus warblers.     

Genetic, morphological, and ecological characterization of a hybrid zone that spans a migratory divide

This study characterizes a hybrid zone that spans a migratory divide between subspecies of the Swainson's thrush (Catharus ustulatus), a long distance migratory songbird, in the Coast Mountains of British Columbia. To assess the potential for a barrier to gene flow between the subspecies, I: (1) analyzed the shape and width of genetic and morphological clines relative to estimates of dispersal distance, (2) assessed the ratio of parental to hybrid genotypes across the hybrid zone, (3) estimated population density across the hybrid zone, and (4) compared the spatial relationship between the hybrid zone and an existing environmental gradient. The results indicate that the hybrid zone is characterized by mostly concordant character clines that are narrow relative to dispersal, the absence of a hybrid swarm, and low population density at the center of the zone. This hybrid zone and additional regions of contact between these subspecies are found on the border between coastal and interior climatic regions throughout the Pacific Northwest. An identified shift in the location, but not the width, of the mtDNA cline relative to the nuclear clines is consistent with asymmetrical hybridization. Neutral diffusion of populations following secondary contact and hybrid superiority within an ecotone are insufficient explanations for the observed patterns. The hypothesis that best fits the data is that the Swainson's thrush hybrid zone is a tension zone maintained by dispersal and ecologically mediated barriers to gene flow.     

Speciation in Progress? A Continuum of Reproductive Isolation in <Emphasis Type="Italic">Drosophila Bipectinata</Emphasis>

Incipient species in the early stages of divergence can provide crucial information about the genetic basis of reproductive isolation and the evolutionary forces that promote speciation. In this report, we describe two subspecies of Drosophila bipectinata that show a continuum of reproductive isolation. Crosses between strains of the same subspecies produce fully fertile offspring. At the same time, each subspecies harbors extensive variation for the degree of reproductive isolation from the other subspecies. The percentage of fertile hybrid males varies from 0 to 90%, depending on the origin of parental strains, indicating that the genes responsible for hybrid sterility are not fixed within either subspecies, or even within local populations. Reproductive isolation is non-transitive, so that the extent of hybrid sterility depends on the particular combination of strains. The two subspecies show little or no evidence of genetic differentiation at three nuclear loci, suggesting that they diverged very recently or continue to experience significant levels of gene flow. A hybrid zone between the two subspecies may exist in New Guinea and Northeastern Australia.     

Varying response of breeding waders to experimental manipulation of their habitat and predators

Many ground-nesting bird species including waders are of conservation concern across Europe. Population declines of Eurasian Curlew Numenius arquata (hereafter Curlew), a globally Near-threatened wader, exemplify this and are driven by low breeding success due to predation, and loss and degradation of breeding habitat. We tested whether the combined delivery of vegetation management and lethal control of Red Foxes Vulpes vulpes and Carrion Crows Corvus corone and Hooded Crows C. cornix improved Curlew nesting success and breeding abundance across UK study sites over five years. On trial sites, vegetation management improved habitat condition and Curlew distributions shifted towards managed areas. However, changes in predator abundances, Curlew nesting success and Curlew breeding abundance did not differ between trial and reference sites. Northern Lapwing Vanellus vanellus numbers increased on trial sites and decreased on reference sites, whilst Common Snipe Gallinago gallinago numbers showed no treatment response. Some of these outcomes were associated with additional environmental variables added to Treatment:Year terms in three-way interactions (changes in fox abundance associated with abundance of non-native gamebirds; changes in Crow abundance associated with gamebirds and woodland area; changes in Curlew nesting success associated with crow abundance), highlighting that outcomes depend on site-specific contexts, and consistent responses to interventions cannot be assumed. The difficulty in increasing Curlew nesting success likely relates to underlying mesopredator densities in the UK, which are known to be high in a European context. Our results suggest that a model of delivery like this study, including management of predators using lethal control, at comparable intensities, is highly unlikely to be effective for Curlew or Snipe within agri-environment schemes, but could be for Northern Lapwing. To make progress on wader recovery it is imperative to understand the underlying drivers of high mesopredator densities and address these through landscape-scale intervention and policy changes.     

Sex chromosome inversions enforce reproductive isolation across an avian hybrid zone

Across hybrid zones, the sex chromosomes are often more strongly differentiated than the autosomes. This is regularly attributed to the greater frequency of reproductive incompatibilities accumulating on sex chromosomes and their exposure in the heterogametic sex. Working within an avian hybrid zone, we explore the possibility that chromosome inversions differentially accumulate on the Z chromosome compared to the autosomes and thereby contribute to Z chromosome differentiation. We analyse the northern Australian hybrid zone between two subspecies of the long‐tailed finch (Poephila acuticauda), first described based on differences in bill colour, using reduced‐representation genomic sequencing for 293 individuals over a 1,530‐km transect. Autosomal differentiation between subspecies is minimal. In contrast, 75% of the Z chromosome is highly differentiated and shows a steep genomic cline, which is displaced 350 km to the west of the cline in bill colour. Differentiation is associated with two or more putative chromosomal inversions, each predominating in one subspecies. If inversions reduce recombination between hybrid incompatibilities, they are selectively favoured and should therefore accumulate in hybrid zones. We argue that this predisposes inversions to differentially accumulate on the Z chromosome. One genomic region affecting bill colour is on the Z, but the main candidates are on chromosome 8. This and the displacement of the bill colour and Z chromosome cline centres suggest that bill colour has not strongly contributed to inversion accumulation. Based on cline width, however, the Z chromosome and bill colour both contribute to reproductive isolation established between this pair of subspecies.     

Identification of 20 polymorphic microsatellite loci in European crow (Corvus corone) from existing passerine loci

The European crow (Corvus corone) occurs in two subspecies (or species) with distinct plumage coloration: the black carrion crow (C. c. corone) and the grey and black hooded crow (C. c. cornix). We tested 42 passerine microsatellite loci for amplification in the European crow and identified 20 loci that were both polymorphic and easy to score. In 50 individuals sampled in the Danish part of the species’ pan‐European hybrid zone, the number of alleles ranged between two and 21. One locus deviated from Hardy–Weinberg equilibrium and had a high estimated null allele frequency. These 20 loci were highly successful in amplifying polymorphic products also in other crow populations and in another Corvidae species, the rook (Corvus frugilegus).     

INCIPIENT SPECIATION DESPITE LITTLE ASSORTATIVE MATING: THE YELLOW‐RUMPED WARBLER HYBRID ZONE

Hybrid zones between recently diverged taxa are natural laboratories for speciation research, allowing us to determine whether there is reproductive isolation between divergent forms and the causes of that isolation. We present a study of a classic avian hybrid zone in North America between two subspecies of the yellow‐rumped warbler (Dendroica coronata). Although previous work has shown very little differentiation in mitochondrial DNA across this hybrid zone, we identified two nuclear loci (one sex‐linked and one autosomal) that show fixed differences across the hybrid zone, in a close concordance with patterns of plumage variation. Temporal stability and limited width of the hybrid zone, along with substantial linkage disequilibrium between these two diagnostic markers in the center of the zone, indicate that there is moderate reproductive isolation between these populations, with an estimated strength of selection maintaining the zone of 18%. Pairing data indicate that assortative mating is either very weak or absent, suggesting that this reproductive isolation is largely due to postmating barriers. Thus, despite extensive hybridization the two forms are distinct evolutionary groups carrying genes for divergent adaptive peaks, and this situation appears relatively stable.     

Divergence in cuticular hydrocarbons between parapatric subspecies of the meadow grasshopper, Chorthippus parallelus (Orthoptera, Acrididae)

Two subspecies of the grasshopper Chorthippus parallelus (Orthoptera; Acrididae) meet and form a hybrid zone in the Pyrenees. Here we investigate whether the subspecies differ in the blend of cuticular hydrocarbons. Such differences may function as chemical signals, being detected via contact chemoreception and used during mate choice as in other insects. Gas chromatography shows that the subspecies have diverged in the composition of their cuticular hydrocarbons. This difference between subspecies exists over and above variation in cuticular hydrocarbons among individual populations. The majority of the difference can be attributed to the relative amounts of a small group of compounds present in the cuticle. The possible consequences of this chemical divergence for mating encounters between the subspecies are discussed.     

The uncharacterized gene 1700093K21Rik and flanking regions are correlated with reproductive isolation in the house mouse,Mus musculus

Reproductive barriers exist between the house mouse subspecies, Mus musculus musculus and M. m. domesticus, members of the Mus musculus species complex, primarily as a result of hybrid male infertility, and a hybrid zone exists where their ranges intersect in Europe. Using single nucleotide polymorphisms (SNPs) diagnostic for the two taxa, the extent of introgression across the genome was previously compared in these hybrid populations. Sixty-nine of 1316 autosomal SNPs exhibited reduced introgression in two hybrid zone transects suggesting maladaptive interactions among certain loci. One of these markers is within a region on chromosome 11 that, in other studies, has been associated with hybrid male sterility of these subspecies. We assessed sequence variation in a 20 Mb region on chromosome 11 flanking this marker, and observed its inclusion within a roughly 150 kb stretch of DNA showing elevated sequence differentiation between the two subspecies. Four genes are associated with this genomic subregion, with two entirely encompassed. One of the two genes, the uncharacterized 1700093K21Rik gene, displays distinguishing features consistent with a potential role in reproductive isolation between these subspecies. Along with its expression specifically within spermatogenic cells, we present various sequence analyses that demonstrate a high rate of molecular evolution of this gene, as well as identify a subspecies amino acid variant resulting in a structural difference. Taken together, the data suggest a role for this gene in reproductive isolation.