Chromatin accessibility, not 5mC methylation covaries with partial dosage compensation in crows.

The evolution of genetic sex determination is often accompanied by degradation of the sex-limited chromosome. Male heterogametic systems have evolved convergent, epigenetic mechanisms restoring the resulting imbalance in gene dosage between diploid autosomes (AA) and the hemizygous sex chromosome (X). Female heterogametic systems (AAf Zf, AAm ZZm) tend to only show partial dosage compensation (0.5 < Zf:AAf < 1) and dosage balance (0.5

Association neurons in the crow telencephalon link visual signs to numerical values.

Many animals can associate signs with numerical values and use these signs in a goal-directed way during task performance. However, the neuronal basis of this semantic association has only rarely been investigated, and so far only in primates. How mechanisms of number associations are implemented in the distinctly evolved brains of other animal taxa such as birds is currently unknown. Here, we explored this semantic number-sign mapping by recording single-neuron activity in the crows' nidopallium caudolaterale (NCL), a brain structure critically involved in avian numerical cognition. Crows were trained to associate visual shapes with varying numbers of items in a number production task. The responses of many NCL neurons during stimulus presentation reflected the numerical values associated with visual shapes in a behaviorally relevant way. Consistent with the crow's better behavioral performance with signs, neuronal representations of numerical values extracted from shapes were more selective compared to those from dot arrays. The existence of number association neurons in crows points to a phylogenetic preadaptation of the brains of cognitively advanced vertebrates to link visual shapes with numerical meaning.

Exogenous and endogenous spatial attention in crows.

Attention describes the ability to selectively process a particular aspect of the environment at the expense of others. Despite the significance of selective processing, the types and scopes of attentional mechanisms in nonprimate species remain underexplored. We trained four carrion crows in Posner spatial cueing tasks using two separate protocols where the attention-capturing cues are shown at different times before target onset at either the same or a different location as the impending target. To probe automatic bottom-up, or exogenous, attention, two naïve crows were tested with a cue that had no predictive value concerning the location of the subsequent target. To examine volitional top-down, or endogenous, attention, the other two crows were tested with the previously learned cues that predicted the impending target location. Comparing the performance for valid (cue and target at same location) and invalid (cue and target at opposing locations) cues in the nonpredictive cue condition showed a transient, mild reaction time advantage signifying exogenous attention. In contrast, there was a strong and long-lasting performance advantage for the valid conditions with predictive cues indicating endogenous attention. Together, these results demonstrate that crows possess two different attention mechanisms (exogenous and endogenous). These findings signify that crows possess a substantial attentional capacity and robust cognitive control over attention allocation.

Conscious Experience of Stimulus Presence and Absence Is Actively Encoded by Neurons in the Crow Brain.

The emergence of consciousness from brain activity constitutes one of the great riddles in biology. It is commonly assumed that only the conscious perception of the presence of a stimulus elicits neuronal activation to signify a "neural correlate of consciousness," whereas the subjective experience of the absence of a stimulus is associated with a neuronal resting state. Here, we demonstrate that the two subjective states "stimulus present" and "stimulus absent" are represented by two specialized neuron populations in crows, corvid birds. We recorded single-neuron activity from the nidopallium caudolaterale of crows trained to report the presence or absence of images presented near the visual threshold. Because of the task design, neuronal activity tracking the conscious "present" versus "absent" percept was dissociated from that involved in planning a motor response. Distinct neuron populations signaled the subjective percepts of "present" and "absent" by increases in activation. The response selectivity of these two neuron populations was similar in strength and time course. This suggests a balanced code for subjective "presence" versus "absence" experiences, which might be beneficial when both conscious states need to be maintained active in the service of goal-directed behavior.

Gaze tracking of large-billed crows (Corvus macrorhynchos) in a motion capture system.

Previous studies often inferred the focus of a bird's attention from its head movements because it provides important clues about their perception and cognition. However, it remains challenging to do so accurately, as the details of how they orient their visual field toward the visual targets remain largely unclear. We thus examined visual field configurations and the visual field use of large-billed crows (Corvus macrorhynchos Wagler 1827). We used an established ophthalmoscopic reflex technique to identify the visual field configuration, including the binocular width and optical axes, as well as the degree of eye movement. A newly established motion capture system was then used to track the head movements of freely moving crows to examine how they oriented their reconstructed visual fields toward attention-getting objects. When visual targets were moving, the crows frequently used their binocular visual fields, particularly around the projection of the beak-tip. When the visual targets stopped moving, crows frequently used non-binocular visual fields, particularly around the regions where their optical axes were found. On such occasions, the crows slightly preferred the right eye. Overall, the visual field use of crows is clearly predictable. Thus, while the untracked eye movements could introduce some level of uncertainty (typically within 15 deg), we demonstrated the feasibility of inferring a crow's attentional focus by 3D tracking of their heads. Our system represents a promising initial step towards establishing gaze tracking methods for studying corvid behavior and cognition.

Hooded crows (Corvus cornix) manufacture objects relative to a mental template.

It was recently found that not only tool-specialized New Caledonian crows, but also Goffin cockatoos can manufacture physical objects in accordance with a mental template. That is, they can emulate features of existing objects when they manufacture new items. Both species spontaneously ripped pieces of card into large strips if they had previously learned that a large template was rewarded, and small strips when they previously learned that a small template was rewarded. Among New Caledonian crows, this cognitive ability was suggested as a potential mechanism underlying the transmission of natural tool designs. Here, we tested for the same ability in another non-specialised tool user-Hooded crows (Corvus cornix). Crows were exposed to pre-made template objects, varying first in colour and then in size, and were rewarded only if they chose pre-made objects that matched the template. In subsequent tests, birds were given the opportunity to manufacture versions of these objects. All three crows ripped paper pieces from the same colour material as the rewarded template, and, crucially, also manufactured objects that were more similar in size to previously rewarded, than unrewarded, templates, despite the birds being rewarded at random in both tests. Therefore, we found the ability to manufacture physical objects relative to a mental template in yet another bird species not specialized in using or making foraging tools in the wild, but with a high level of brain and cognitive development.

Fitness benefits of alternated chick provisioning in cooperatively breeding carrion crows.

In most bird species, parents raise offspring cooperatively. In some cases, this cooperation extends to helpers-at-the-nest who assist the breeders with a range of tasks. While cooperative food provisioning might merely arise incidentally, as a result of the efforts of carers that act independently from each other, recent studies suggest that birds may coordinate by taking turns in visiting the nest. However, evidence that such coordination emerges because individuals actively respond to each other's behaviour is controversial, and the potential benefits of carers' alternation remain unknown. We addressed this knowledge gap by analysing a multiyear dataset for cooperatively breeding carrion crows Corvus corone, comprising 8693 nest visits across 50 groups. Our results reveal that turn-taking does occur in this species and that all group members, regardless of their sex and social role (breeder/helper), tend to alternate at the nest with other carers rather than to make repeat visits. Importantly, we found that the body mass of nestlings increased significantly with the degree of carers' alternation, possibly because well-coordinated groups provided food at more regular intervals. Using earlier monitoring data, the observed increase in body mass is predicted to substantially boost postfledging survival rates. Our analyses demonstrate that alternation in nestling provisioning has measurable fitness benefits in this study system. This raises the possibility that cooperatively breeding carrion crows, as well as other bird species with similarly coordinated brood provisioning, exhibit specialized behavioural strategies that enable effective alternation.

En la mayoría de las especies de aves, los padres cooperan entre sí en el cuidado de sus crías. En algunos casos, la cooperación se extiende a individuos ayudantes que asisten a los reproductores en diferentes tareas. El aprovisionamiento cooperativo de las crías puede darse simplemente como resultado del esfuerzo de cuidadores que actúan de forma independiente, pero estudios recientes sugieren que las aves pueden coordinarse, tomando turnos a la hora de visitar el nido. Sin embargo, las evidencias de que dicha coordinación ocurre porque los individuos realmente responden al comportamiento de los demás es controvertida, y los potenciales beneficios de la alternancia al nido son desconocidos. Nosotros abordamos estos aspectos analizando una base de datos plurianual en la corneja negra Corvus corone, que incluye 8693 visitas al nido en 50 grupos. Nuestros resultados revelan que se produce toma de turnos en esta especie y que todos los miembros del grupo, independientemente del sexo y del estatus social (reproductor / ayudante), tienden a alternarse al nido con otros cuidadores, en vez de repetir visitas. Más importante aún, encontramos que la masa corporal de los polluelos aumentaba de forma significativa al aumentar el grado de coordinación de los cuidadores, posiblemente porque los grupos mejor coordinados aprovisionaban las crías a intervalos más regulares. Basándonos en datos anteriores, pudimos calcular que el incremento observado en masa corporal predice un aumento sustancial de la tasa de supervivencia de los volanderos. Nuestros análisis demuestran, por lo tanto, que la alternancia al nido tiene beneficios medibles en eficacia biológica. Esto conlleva la posibilidad de que las cornejas negras cooperativas, al igual que otras especies de aves con aprovisionamiento coordinado de las crías, exhiban estrategias comportamentales especializadas que permitan una eficiente alternancia al nido.

Reduction of genetic diversity in 'Alala (Hawaiian crow; Corvus hawaiiensis) between the late 1800s and the late 1900s.

Genetic and genomic data are increasingly used to aid conservation management of endangered species by providing insights into evolutionary histories, factors associated with extinction risks, and potential for future adaptation. For the 'Alala, or Hawaiian crow (Corvus hawaiiensis), genetic concerns include negative correlations between inbreeding and hatching success. However, it is unclear if low genetic diversity and inbreeding depression are consequences of a historical population bottleneck, or if 'Alala had historically low genetic diversity that predated human influence, perhaps as a result of earlier declines or founding events. In this study, we applied a hybridization-based sequence capture to generate a genome-wide single nucleotide polymorphism (SNP) dataset for comparing historical specimens collected in the 1890s, when 'Alala were more numerous, to samples taken between 1973 and 1998, when 'Alala population densities were near the lowest documented levels in the wild, prior to all individuals being collected for captive rearing. We found low genome-wide diversity in both sample groups, however, the modern sample group (1973 to 1998 cohort) exhibited relatively fewer polymorphic alleles, a lower proportion of polymorphic loci, and lower observed heterozygosity, consistent with a population decline and potential bottleneck effects. These results combined with a current low population size highlight the importance of continued efforts by conservation managers to mitigate inbreeding and maintain founder representation to preserve what genetic diversity remains.

DNA barcoding identifies cryptic animal tool materials.

Some animals fashion tools or constructions out of plant materials to aid foraging, reproduction, self-maintenance, or protection. Their choice of raw materials can affect the structure and properties of the resulting artifacts, with considerable fitness consequences. Documenting animals' material preferences is challenging, however, as manufacture behavior is often difficult to observe directly, and materials may be processed so heavily that they lack identifying features. Here, we use DNA barcoding to identify, from just a few recovered tool specimens, the plant species New Caledonian crows (Corvus moneduloides) use for crafting elaborate hooked stick tools in one of our long-term study populations. The method succeeded where extensive fieldwork using an array of conventional approaches-including targeted observations, camera traps, radio-tracking, bird-mounted video cameras, and behavioral experiments with wild and temporarily captive subjects-had failed. We believe that DNA barcoding will prove useful for investigating many other tool and construction behaviors, helping to unlock significant research potential across a wide range of study systems.

Competition and generalization impede cultural formation in wild jackdaws.

Animal cultures have now been demonstrated experimentally in diverse taxa from flies to great apes. However, experiments commonly use tasks with unrestricted access to equal pay-offs and innovations seeded by demonstrators who are trained to exhibit strong preferences. Such conditions may not reflect those typically found in nature. For example, the learned preferences of natural innovators may be weaker, while competition for depleting resources can favour switching between strategies and generalizing from past experience. Here we show that in experiments where wild jackdaws (Corvus monedula) can freely discover depleting supplies of novel foods, generalization has a powerful effect on learning, allowing individuals to exploit multiple new opportunities through both social and individual learning. Further, in contrast to studies with trained demonstrators, individuals that were first to innovate showed weak preferences. As a consequence, many individuals ate all available novel foods, displaying no strong preference and no group-level culture emerged. Individuals followed a 'learn from adults' strategy, but other demographic factors played a minimal role in shaping social transmission. These results demonstrate the importance of generalization in allowing animals to exploit new opportunities and highlight how natural competitive dynamics may impede the formation of culture.

Satellite DNA evolution in Corvoidea inferred from short and long reads.

Satellite DNA (satDNA) is a fast-evolving portion of eukaryotic genomes. The homogeneous and repetitive nature of such satDNA causes problems during the assembly of genomes, and therefore it is still difficult to study it in detail in nonmodel organisms as well as across broad evolutionary timescales. Here, we combined the use of short- and long-read data to explore the diversity and evolution of satDNA between individuals of the same species and between genera of birds spanning ~40 millions of years of bird evolution using birds-of-paradise (Paradisaeidae) and crow (Corvus) species. These avian species highlighted the presence of a GC-rich Corvoidea satellitome composed of 61 satellite families and provided a set of candidate satDNA monomers for being centromeric on the basis of length, abundance, homogeneity and transcription. Surprisingly, we found that the satDNA of crow species rapidly diverged between closely related species while the satDNA appeared more similar between birds-of-paradise species belonging to different genera.

Ectoparasite presence and brood size manipulation interact to accelerate telomere shortening in nestling jackdaws.

Early-life conditions impact fitness, but whether the combined effect of extrinsic stressors is additive or synergistic is not well known. This is a major knowledge gap because exposure to multiple stressors is frequent. Telomere dynamics may be instrumental when testing how stressors interact because many factors affect telomere shortening, and telomere shortening predicts survival. We evaluated the effect of manipulated brood size and natural infestation by the carnid fly Carnus hemapterus on nestling growth and telomere shortening of wild jackdaws (Corvus monedula). Telomere length, measured in blood using TRF, shortened on average by 264 bp, and on average, Carnus infection induced more telomere shortening. Further analyses showed that in enlarged broods, nestlings' telomeres shortened more when parasitized, while in reduced broods there was no effect of infection on telomere shortening. We conclude that there is a synergistic effect of number of siblings and Carnus infection on telomere shortening rate: blood-sucking parasites may negatively impact telomeres by increasing cell proliferation and/or physiological stress, and coping with infection may be less successful in enlarged broods with increased sibling competition. Larger nestlings had shorter telomeres independent of age, brood manipulation or infection. Growth was independent of infestation but in enlarged broods, nestlings were lighter at fledging. Our findings indicate that (i) evaluating consequences of early-life environmental conditions in isolation may not yield a full picture due to synergistic effects, and (ii) effects of environmental conditions may be cryptic, for example, on telomeres, with fitness consequences expressed beyond the temporal framework of the study.

Increased genetic diversity and immigration after West Nile virus emergence in American crows: No evidence for a genetic bottleneck.

Infectious diseases can cause steep declines in wildlife populations, leading to changes in genetic diversity that may affect the susceptibility of individuals to infection and the overall resilience of populations to pathogen outbreaks. Here, we examine evidence for a genetic bottleneck in a population of American crows (Corvus brachyrhynchos) before and after the emergence of West Nile virus (WNV). More than 50% of marked birds in this population were lost over the 2-year period of the epizootic, representing a 10-fold increase in adult mortality. Using analyses of single-nucleotide polymorphisms (SNPs) and microsatellite markers, we tested for evidence of a genetic bottleneck and compared levels of inbreeding and immigration in the pre- and post-WNV populations. Counter to expectations, genetic diversity (allelic diversity and the number of new alleles) increased after WNV emergence. This was likely due to increases in immigration, as the estimated membership coefficients were lower in the post-WNV population. Simultaneously, however, the frequency of inbreeding appeared to increase: Mean inbreeding coefficients were higher among SNP markers, and heterozygosity-heterozygosity correlations were stronger among microsatellite markers, in the post-WNV population. These results indicate that loss of genetic diversity at the population level is not an inevitable consequence of a population decline, particularly in the presence of gene flow. The changes observed in post-WNV crows could have very different implications for their response to future pathogen risks, potentially making the population as a whole more resilient to a changing pathogen community, while increasing the frequency of inbred individuals with elevated susceptibility to disease.

Highly pathogenic avian influenza (H5N1) infection in crows through ingestion of infected crow carcasses.

The present study was aimed to investigate the role of cannibalism in transmission of H5N1 avian influenza virus to house crows (Corvus splendens). Four crows were intranasally inoculated with 108.0 EID50 (A/crow/India/01CA249/2021) H5N1 highly pathogenic avian influenza (HPAI) virus and were observed for 14 days for any overt signs of illness. Two of the infected crows showed signs of wing paralysis, incoordination, and torticollis. For cannibalism experiment, two crows showing clinical signs were euthanized on 14th day post-infection (dpi) and were kept in the isolator and four naïve healthy crows were introduced along with the euthanized crows. The viscera from the infected carcasses were eaten by all the four crows. Oropharyngeal and cloacal swabs were collected up to 14 days to assess virus excretion. All four crows showed clinical signs viz., dullness, reluctance to move with ruffled feathers on 6th day post cannibalism along with neurological signs including incoordination and paralysis of the wings. All the crows gradually recovered after showing clinical signs and were euthanized on 21st day of observation period. Virus excretion was observed from 3rd to 11th day post cannibalism through both oropharyngeal and cloacal routes with maximum shedding through oropharyngeal route. The virus was isolated from lungs and trachea of one the infected crows at 21st day after euthanasia. All the four crows seroconverted against H5N1 virus infection at 14th day post cannibalism. Our study confirms the transmission of H5N1 virus in crows through cannibalism and highlights how H5N1 virus might circulate in a crow colony once they become infected.

Numerical Representation for Action in Crows Obeys the Weber-Fechner Law.

The psychophysical laws governing the judgment of perceived numbers of objects or events, called the number sense, have been studied in detail. However, the behavioral principles of equally important numerical representations for action are largely unexplored in both humans and animals. We trained two male carrion crows (Corvus corone) to judge numerical values of instruction stimuli from one to five and to flexibly perform a matching number of pecks. Our quantitative analysis of the crows' number production performance shows the same behavioral regularities that have previously been demonstrated for the judgment of sensory numerosity, such as the numerical distance effect, the numerical magnitude effect, and the logarithmical compression of the number line. The presence of these psychophysical phenomena in crows producing number of pecks suggests a unified sensorimotor number representation system underlying the judgment of the number of external stimuli and internally generated actions.

A fundamental propulsive mechanism employed by swimmers and flyers throughout the animal kingdom.

Even casual observations of a crow in flight or a shark swimming demonstrate that animal propulsive structures bend in patterned sequences during movement. Detailed engineering studies using controlled models in combination with analysis of flows left in the wakes of moving animals or objects have largely confirmed that flexibility can confer speed and efficiency advantages. These studies have generally focused on the material properties of propulsive structures (propulsors). However, recent developments provide a different perspective on the operation of nature's flexible propulsors, which we consider in this Commentary. First, we discuss how comparative animal mechanics have demonstrated that natural propulsors constructed with very different material properties bend with remarkably similar kinematic patterns. This suggests that ordering principles beyond basic material properties govern natural propulsor bending. Second, we consider advances in hydrodynamic measurements demonstrating suction forces that dramatically enhance overall thrust produced by natural bending patterns. This is a previously unrecognized source of thrust production at bending surfaces that may dominate total thrust production. Together, these advances provide a new mechanistic perspective on bending by animal propulsors operating in fluids - either water or air. This shift in perspective offers new opportunities for understanding animal motion as well as new avenues for investigation into engineered designs of vehicles operating in fluids.

Crows protect visual working memory against interference.

Working memory, the ability to actively maintain and manipulate information across time, is key to intelligent behavior. Because of the limited capacity of working memory, relevant information needs to be protected against distracting representations. Whether birds can resist distractors and safeguard memorized relevant information is unclear. We trained carrion crows in a delayed match-to-sample task to memorize an image while resisting other, interfering stimuli. We found that the repetition of the sample stimulus during the memory delay improved performance accuracy and accelerated reaction time relative to a reference condition with a neutral interfering stimulus. In contrast, the presentation of the image that constituted the subsequent non-match test stimulus mildly weakened performance. However, the crows' robust performance in this most demanding distractor condition indicates that sample information was actively protected from being overwritten by the distractor. These data show that crows can cognitively control and safeguard behaviorally relevant working memory contents.

Are you better than me? Social comparisons in carrion crows (Corvus corone).

Comparing oneself to others is a key process in humans that allows individuals to gauge their performances and abilities and thus develop and calibrate their self-image. Little is known about its evolutionary foundations. A key feature of social comparison is the sensitivity to other individuals' performance. Recent studies on primates produced equivocal results, leading us to distinguish between a 'strong' variant of the social comparison hypothesis formulated for humans and a 'weak' variant found in non-human primates that would comprise some elements of human social comparison. Here, we focus on corvids that are distantly related to primates and renowned for their socio-cognitive skills. We were interested in whether crows' task performances were influenced (i) by the presence of a conspecific co-actor performing the same discrimination task and (ii) by the simulated acoustic cues of a putative co-actor performing better or worse than themselves. Crows reached a learning criterion quicker when tested simultaneously as compared to when tested alone, indicating a facilitating effect of social context. The performance of a putative co-actor influenced their performance: crows were better at discriminating familiar images when their co-actor was better than they were. Standard extremity (how pronounced the difference was between the performance of the subject and that of the co-actor), and category membership (affiliation status and sex), of the putative co-actors had no effect on their performance. Our findings are in line with the 'weak' variant of social comparison and indicate that elements of human social comparison can be found outside of primates.

Differential responses to con- and allospecific visual cues in juvenile ravens (Corvus corax): the ontogeny of gaze following and social predictions.

Gaze following refers to the ability to co-orient with others' gaze directions. Ontogenetic studies on gaze following in animals have predominantly used human experimenters as demonstrators. It is, however, likely that developing animals are initially more attuned to individuals from their own species, which might result in differences in the ontogenetic onset of gaze following with human and conspecific demonstrators. "Checking back" is a signature behaviour in the gaze following repertoires of humans, apes, and some Old world monkeys. It is commonly interpreted as a representation of the referentiality of gaze and is thus diagnostic of social predictions. Recently, "checking back" has been discovered in four avian species, suggesting a shared skill among birds. To investigate effects of con- and allospecific demonstrators on gaze following responses, we studied visual co-orientations of four hand-raised juvenile common ravens (Corvus corax) with human and conspecific gaze cues. Moreover, we for the first time investigated "checking back" in ravens and compared the effects of con- and allospecific demonstrators on this behaviour. Ravens followed human and conspecific gaze with no apparent differences in ontogenetic onset, but after significantly longer latencies with human demonstrators. Subjects moreover already checked back at 30 days old and did so significantly more often with conspecific demonstrators. Our findings suggest differences in processing speed and social predictions of human and conspecific gazes, indicating an underlying neurocognitive mechanism attuned to social information gathering from conspecifics. We propose more studies using conspecific demonstrators to reveal the full gaze following potential of a species.

Oscillatory extraction behaviour suggests functional attributes of crows' hooked-stick tools.

New Caledonian crows are the only nonhuman animals known to craft hooked-sticks for use in foraging. Since their first description over 25 years ago, researchers have been unable to provide a detailed account of how these complex tools function in natural probe sites. Using close-up video footage, we document how a New Caledonian crow operated a hooked-stick to extract a large tree weta from a chamber in a tree trunk. The extraction technique had two distinct, separate components: (1) simultaneous oscillating head rotation and reciprocating bill action, and (2) measured pulling with the tool. Analysis of this first detailed field observation of hooked-stick use suggests a link between hooked-stick tool characteristics, functionality and skilled manipulation in natural prey extraction by these technological birds. Our findings also provide a rare, if not novel, example of tool-associated oscillatory manipulation in nonhuman animals.