Differential calls by Carrion and Hooded Crows (Corvus corone corone and C. c. cornix) in the Alpine hybrid zone

Spectrograms of calls given by Carrion and Hooded Crows in the Italian Alpine hybrid zone have been tested for differences between the two subspecies by univariate and multivariate techniques. Vocalizations of the two Crows are similar and composed of 2–15 caws, each usually made up of two harmonics, separated by short intervals. However, one-way ANOVA and discriminant analysis showed that they were differentiated with respect both to temporal and frequency parameters. We suggest that differences in vocalizations reflect mostly the phylogenetic divergence between Hooded and Carrion Crows. The results are in keeping with previous ecological, morphological and molecular data and would confirm that the present designation of the two Crows as distinct subspecies is correct. Differences in vocalizations, together with others regarding ecology and behaviour, may result in assortative mating and homotypic flocking and, therefore, may contribute to reinforcement of the reproductive isolation between the Carrion and the Hooded Crow in the Alpine hybrid zone.     

A SOCIO-ECOLOGY OF MAGPIES PICA PICA

The behaviour and ecology of unmarked Magpies Pica pica living in the Cotswolds were investigated over 18 months. The spacing of nests was significantly more regular than random and breeding pairs defended territories throughout the year. In winter pairs occasionally left territories to feed with non-breeding birds which flocked throughout the year.
Magpies in flocks had a higher averiage feeding rate than birds alone or in pairs. Aggression rates in flocks increased as birds fed closer together and as food patches became richer.
Agonistic interactions between Crows Corvus corone and Magpies were common; Crows frequently dominated Magpies in disputes over food, and Crows also ate Magpie eggs and young. Single Magpies were most often chased by Crows while Magpies in flocks were able to feed longer in the presence of Crows.
Differences in fledgling success among breeding pairs were related to the location of nests; Magpie pairs breeding near Crow nests suffered higher rates of agonistic encounters with Crows and also produced fewer fledglings than did Magpies breeding at greater distances from Crow nests. Magpie nests were located nearer human habitations than were Crow nests.     

Individual and social factors affecting the ability of American crows to solve and master a string pulling task

Crows and other birds in the family Corvidae regularly share information to learn the identity and whereabouts of dangerous predators, but can they use social learning to solve a novel task for a food reward? Here, we examined the factors affecting the ability of 27 wild-caught American crows to solve a common string pulling task in a laboratory setting. We split crows into two groups; one group was given the task after repeatedly observing a conspecific model the solution and the other solved in the absence of conspecific models. We recorded the crows’ estimated age, sex, size, body condition, level of nervousness, and brain volume using DICOM images from a CT scan. Although none of these variables were statistically significant, crows without a conspecific model and large brain volumes consistently mastered the task in the minimum number of days, whereas those with conspecific models and smaller brain volumes required varying and sometimes a substantial number of days to master the task. We found indirect evidence that body condition might also be important for motivating crows to solve the task. Crows with conspecific models were no more likely to initially solve the task than those working the puzzle without social information, but those that mastered the task usually copied the method most frequently demonstrated by their knowledgeable neighbors. These findings suggest that brain volume and possibly body condition may be factors in learning new tasks and that crows can use social learning to refine their ability to obtain a novel food source, although they must initially learn to access it themselves.     

Co-breeding involving herons and a potential egg predator,the Indian House Crow (Corvus splendens), in Peninsular India

Association between species may strengthen the fitness of the species involved It is not rare that avian species associate on the breeding and feeding grounds. However, a species associated with a potential egg predator is less common. In this study, a synchronized breeding of Indian House Crow (Corvus splendens) and breeding Indian Pond Herons (Ardeola grayii) in urban conditions is reported. Both the crow abundance and the crow nest abundance increased with the number of heronry nests on sites. Crows were mostly observed when flying over or when resting nearby, but they also attempted egg predation from heronry nests. Crows also used the heronry sites for collecting nesting resources, such as twigs, scavenging dead chicks and for stealing the food brought to feed the heronry chicks. A dearth of suitable nesting places and provisions in an urban environment may be the reason why these birds share nesting trees. Vigilant breeding crows, despite their ability to depredate heron nests, may be more beneficial to herons as they are known to mob and distract heron predators, but a full cost-benefit analysis needs to be undertaken.     

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.     

Complex cognition and behavioural innovation in New Caledonian crows

Apes, corvids and parrots all show high rates of behavioural innovation in the wild. However, it is unclear whether this innovative behaviour is underpinned by cognition more complex than simple learning mechanisms. To investigate this question we presented New Caledonian crows with a novel three-stage metatool problem. The task involved three distinct stages: (i) obtaining a short stick by pulling up a string, (ii) using the short stick as a metatool to extract a long stick from a toolbox, and finally (iii) using the long stick to extract food from a hole. Crows with previous experience of the behaviours in stages 1–3 linked them into a novel sequence to solve the problem on the first trial. Crows with experience of only using string and tools to access food also successfully solved the problem. This innovative use of established behaviours in novel contexts was not based on resurgence, chaining and conditional reinforcement. Instead, the performance was consistent with the transfer of an abstract, causal rule: ‘out-of-reach objects can be accessed using a tool’. This suggests that high innovation rates in the wild may reflect complex cognitive abilities that supplement basic learning mechanisms.     

Injury and paternity loss in cooperatively breeding American Crows

ABSTRACT Although survivable injuries have been documented in several species of birds, little is known about the social and reproductive consequences of such injuries. We examined relationships between injuries and group composition, reproductive output, and paternity of male breeders in 27 family groups of American Crows (Corvus brachyrhynchos) using 96 broods sampled from 2004 to 2009. Injuries and permanent disfigurement of wings or feet were sustained by 25.9% (7/27) of dominant male breeders. Injured male breeders had smaller broods (2.13 vs. 3.38 nestlings per brood for injured vs. uninjured breeders), lost more paternity (48% vs. 13% extrapair young per brood), and had more adult male auxiliaries in their groups (1.63 vs. 0.65 auxiliaries per group) than uninjured birds. Some of the variation in reproductive partitioning, extrapair paternity rates, and group composition among American Crow family groups can therefore be explained by injuries to male breeders. These results suggest that injuries could account for some of the unexplained variation in the rates and distribution of extrapair paternity across populations and taxa, as well as some of the variation in reproductive skew among cooperatively breeding birds.     

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.

     

An island-wide predator manipulation reveals immediate and long-lasting matching of risk by prey

Anti-predator behaviour affects prey population dynamics, mediates cascading effects in food webs and influences the likelihood of rapid extinctions. Predator manipulations in natural settings provide a rare opportunity to understand how prey anti-predator behaviour is affected by large-scale changes in predators. Here, we couple a long-term, island-wide manipulation of an important rodent predator, the island fox (Urocyon littoralis), with nearly 6 years of measurements on foraging by deer mice (Peromyscus maniculatus) to provide unequivocal evidence that prey closely match their foraging behaviour to the number of fox predators present on the island. Peromyscus maniculatus foraging among exposed and sheltered microhabitats (a measure of aversion to predation risk) closely tracked fox density, but the nature of this effect depended upon nightly environmental conditions known to affect rodent susceptibility to predators. These effects could not be explained by changes in density of deer mice over time. Our work reveals that prey in natural settings are cognizant of the dynamic nature of their predators over timescales that span many years, and that predator removals spanning many generations of prey do not result in a loss of anti-predator behaviour.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.     

Hook tool manufacture in New Caledonian crows: behavioural variation and the influence of raw materials

Background

New Caledonian crows use a range of foraging tools, and are the only non-human species known to craft hooks. Based on a small number of observations, their manufacture of hooked stick tools has previously been described as a complex, multi-stage process. Tool behaviour is shaped by genetic predispositions, individual and social learning, and/or ecological influences, but disentangling the relative contributions of these factors remains a major research challenge. The properties of raw materials are an obvious, but largely overlooked, source of variation in tool-manufacture behaviour. We conducted experiments with wild-caught New Caledonian crows, to assess variation in their hooked stick tool making, and to investigate how raw-material properties affect the manufacture process.

Results

In Experiment 1, we showed that New Caledonian crows’ manufacture of hooked stick tools can be much more variable than previously thought (85 tools by 18 subjects), and can involve two newly-discovered behaviours: ‘pulling’ for detaching stems and bending of the tool shaft. Crows’ tool manufactures varied significantly: in the number of different action types employed; in the time spent processing the hook and bending the tool shaft; and in the structure of processing sequences. In Experiment 2, we examined the interaction of crows with raw materials of different properties, using a novel paradigm that enabled us to determine subjects’ rank-ordered preferences (42 tools by 7 subjects). Plant properties influenced: the order in which crows selected stems; whether a hooked tool was manufactured; the time required to release a basic tool; and, possibly, the release technique, the number of behavioural actions, and aspects of processing behaviour. Results from Experiment 2 suggested that at least part of the natural behavioural variation observed in Experiment 1 is due to the effect of raw-material properties.

Conclusions

Our discovery of novel manufacture behaviours indicates a plausible scenario for the evolutionary origins, and gradual refinement, of New Caledonian crows’ hooked stick tool making. Furthermore, our experimental demonstration of a link between raw-material properties and aspects of tool manufacture provides an alternative hypothesis for explaining regional differences in tool behaviours observed in New Caledonian crows, and some primate species.

     

Investigating responses to control: a comparison of common myna behaviour across areas of high and low trapping pressure

Harvesting of wildlife by humans is commonly known to impact target species’ demography, life history and behaviour. Yet in the context of invasive and pest species control, behavioural shifts have received very little attention. The introduced common myna, Acridotheres tristis, is undergoing intensive control efforts in some areas of its Australian distribution. In order to investigate whether myna populations respond to harvesting, we compared the behaviour of free-ranging common mynas in areas of high and low trapping pressure. Behavioural analyses revealed that mynas in high  trapping risk areas stayed closer to refuges, tended to form smaller groups, and were overall less detectable. Behavioural differences between high and lowly trapped areas were not attributable to variation in population density. Overall, these results are consistent with the hypothesis that common mynas have the potential to adjust their behaviour in response to heightened anthropic risk in the environment. Behavioural shifts in invasive alien species could modify their ecological impact and may interfere not only with the effectiveness of control measures, but also with how their effectiveness is assessed. Future research should aim to isolate behavioural mechanisms underpinning compensatory responses to control so that any potential effects can be mitigated.     

Tocopherols and tocotrienols in membranes: a critical review

The familiar role of tocols (tocopherols and tocotrienols) as lipid-soluble chain-terminating inhibitors of lipid peroxidation is currently in the midst of a reinterpretation. New biological activities have been described for tocols that apparently are not dependent on their well-established antioxidant behaviour. These activities could well be real, but there remain large gaps in our understanding of the behaviour of tocols in membranes, especially when it comes to the alpha-, beta-, gamma-, delta-chroman methylation patterns and the seemingly special nature of tocotrienols. It is inappropriate to make conclusions and develop models based on in vivo (or cell culture) results with reference to in vitro measurements of antioxidant activity. When present in biological membranes, tocols will experience a large variation in the local composition of phospholipids and the presence of neutral lipids such as cholesterol, both of which would be expected to change the efficiency of antioxidant action. It is likely that tocols are not homogeneously dispersed in a membrane, but it is still not known whether any specific combination of lipid head group and acyl chains are conferred special protection from peroxidation, nor do we currently appreciate the structural role that tocols play in membranes. Tocols may enhance curvature stress or counteract similar stresses generated by other lipids such as lysolipids. This review will outline what is known about the location and behaviour of tocols in phospholipid bilayers. We will draw mainly from the biophysical literature, but will attempt to extend the discussion to biologically relevant phenomena when appropriate. We hope that it will assist researchers when designing new experiments and when critically assessing the results, in turn providing a more thorough understanding of the biochemistry of tocols.     

The evolutionary origins and ecological context of tool use in New Caledonian crows

New Caledonian (NC) crows Corvus moneduloides are the most prolific avian tool users. In the wild, they use at least three distinct tool types to extract invertebrate prey from deadwood and vegetation, with some of their tools requiring complex manufacture, modification and/or deployment. Experiments with captive-bred, hand-raised NC crows have demonstrated that the species has a strong genetic predisposition for basic tool use and manufacture, suggesting that this behaviour is an evolved adaptation. This view is supported by recent stable-isotope analyses of the diets of wild crows, which revealed that tool use provides access to highly profitable hidden prey, with preliminary data indicating that parents preferentially feed their offspring with tool-derived food. Building on this work, our review examines the possible evolutionary origins of these birds’ remarkable tool-use behaviour. Whilst robust comparative analyses are impossible, given the phylogenetic rarity of animal tool use, our examination of a wide range of circumstantial evidence enables a first attempt at reconstructing a plausible evolutionary scenario. We suggest that a common ancestor of NC crows, originating from a (probably) non-tool-using South-East Asian or Australasian crow population, colonised New Caledonia after its last emersion several million years ago. The presence of profitable but out-of-reach food, in combination with a lack of direct competition for these resources, resulted in a vacant woodpecker-like niche. Crows may have possessed certain behavioural and/or morphological features upon their arrival that predisposed them to express tool-use rather than specialised prey-excavation behaviour, although it is possible that woodpecker-like foraging preceded tool use. Low levels of predation risk may have further facilitated tool-use behaviour, by allowing greater expenditure of time and energy on object interaction and exploration, as well as the evolution of a ‘slow’ life-history, in which prolonged juvenile development enables acquisition of complex behaviours. Intriguingly, humans may well have influenced the evolution of at least some of the species’ tool-oriented behaviours, via their possible introduction of candlenut trees together with the beetle larvae that infest them. Research on NC crows’ tool-use behaviour in its full ecological context is still in its infancy, and we expect that, as more evidence accumulates, some of our assumptions and predictions will be proved wrong. However, it is clear from our analysis of existing work, and the development of some original ideas, that the unusual evolutionary trajectory of NC crows is probably the consequence of an intricate constellation of interplaying factors.     

An analysis of genetic differentiation among assortatively mating Drosophila melanogaster in Zimbabwe

African Drosophila melanogaster populations, and those from Zimbabwe in particular, have attracted much interest recently. African flies differ genetically from 'cosmopolitan' populations and were found to exhibit discriminative mating behaviour against individuals from 'cosmopolitan' populations. It has therefore been proposed that Zimbabwean and some other African populations are in an 'incipient stage of speciation'. However, whether the mating behaviour is an effective barrier against gene flow from other populations, and whether intra-population genetic differentiation has already evolved in sympatry is not known. Here, we took a population-based approach to test whether the well-characterized mating type differences have resulted in a genome-wide differentiation at the population level. Using 122 polymorphic microsatellite loci mapping to the third chromosome, we demonstrate a significant genetic differentiation between Zimbabwean flies differing in their mating behaviour. We also provide evidence to suggest that this difference is unlikely to be attributable to population structure within Zimbabwe. However, the analysis of individual microsatellite loci did not indicate more loci differentiating these two groups than expected by chance. Our data suggest that the 'Z'-'M' mating behaviour is strong enough to result in a small but significant genetic differentiation. Thus, future studies based on a larger population sample of flies characterized for their mating behaviour and using more markers are expected to provide more information on the genetic basis of the mating traits in the Zimbabwe flies.     

Brains, tools, innovation and biogeography in crows and ravens

ABSTRACT: BACKGROUND: Crows and ravens (Passeriformes: Corvus) are large-brained birds with enhanced cognitive abilities relative to other birds. They are among the few non-hominid organisms on Earth to be considered intelligent and well-known examples exist of several crow species having evolved innovative strategies and even use of tools in their search for food. The 40 Corvus species have also been successful dispersers and are distributed on most continents and in remote archipelagos. RESULTS: This study presents the first molecular phylogeny including all species and a number of subspecies within the genus Corvus. We date the phylogeny and determine ancestral areas to investigate historical biogeographical patterns of the crows. Additionally, we use data on brain size and a large database on innovative behaviour and tool use to test whether brain size (i) explains innovative behaviour and success in applying tools when foraging and (ii) has some correlative role in the success of colonization of islands. Our results demonstrate that crows originated in the Palaearctic in the Miocene from where they dispersed to North America and the Caribbean, Africa and Australasia. We find that relative brain size alone does not explain tool use, innovative feeding strategies and dispersal success within crows. CONCLUSIONS: Our study supports monophyly of the genus Corvus and further demonstrates the direction and timing of colonization from the area of origin in the Palaearctic to other continents and archipelagos. The Caribbean was probably colonized from North America, although some North American ancestor may have gone extinct, and the Pacific was colonized multiple times from Asia and Australia. We did not find a correlation between relative brain size, tool use, innovative feeding strategies and dispersal success. Hence, we propose that all crows and ravens have relatively large brains compared to other birds and thus the potential to be innovative if conditions and circumstances are right.     

Eye morphology and visual acuity in the pipevine swallowtail (Battus philenor) studied with a new method of measuring interommatidial angles

Because of the important role sensory systems play in the behaviour of animals, information on sensory capabilities is of great value to behavioural ecologists in the development of hypotheses to explain behaviour. In compound eyes, interommatidial angles are a key determinant of visual acuity but methods for measuring these angles are often demanding and limited to live animals with a pseudopupil. Here we present a new technique for measuring interommatidial angles that is less demanding in terms of technology than other techniques but still accurate. It allows measurements in eyes without a pseudopupil such as dark eyes or even museum specimens. We call this technique the radius of curvature estimation (RCE) method. We describe RCE and validate the method by comparing results from RCE with those from pseudopupil analysis for the butterfly Asterocampa leilia. As an application of RCE we measure the eyes of the butterfly Battus philenor, a species whose visually guided behaviour is well known but whose eye structure and visual acuity are unknown. We discuss the results of the eye morphology in B. philenor in relation to their behaviour and ecology. We contend that RCE fills a gap in the repertoire of techniques available to study peripheral determinants of spatial resolution in compound eyes, because it can be applied on species with dark eyes. RCE then opens up for sampling a larger number of specimens, which, in combination with being able to use museum specimens, makes it possible to quantitatively test ecologically and evolutionarily driven hypotheses about vision in animals in a new way.     

Wild female baboons bias their social behaviour towards paternal half-sisters

Adult female cercopithecines have long been known to bias their social behaviour towards close maternal kin. However, much less is understood about the behaviour of paternal kin, especially in wild populations. Here, we show that wild adult female baboons bias their affiliative behaviour towards their adult paternal half-sisters in the same manner and to the same extent that they bias their behaviour towards adult maternal half-sisters. Females appear to rely heavily on social familiarity as a means of biasing their behaviour towards paternal half-sisters, but may use phenotype matching as well.