On the evolutionary and ontogenetic origins of tool‐oriented behaviour in New Caledonian crows (Corvus moneduloides)

New Caledonian crows (Corvus moneduloides) are prolific tool users in captivity and in the wild, and have an inherited predisposition to express tool‐oriented behaviours. To further understand the evolution and development of tool use, we compared the development of object manipulation in New Caledonian crows and common ravens (Corvus corax), which do not routinely use tools. We found striking qualitative similarities in the ontogeny of tool‐oriented behaviour in New Caledonian crows and food‐caching behaviour in ravens. Given that the common ancestor of New Caledonian crows and ravens was almost certainly a caching species, we therefore propose that the basic action patterns for tool use in New Caledonian crows may have their evolutionary origins in caching behaviour. Noncombinatorial object manipulations had similar frequencies in the two species. However, frequencies of object combinations that are precursors to functional behaviour increased in New Caledonian crows and decreased in ravens throughout the study period, ending 6 weeks post‐fledging. These quantitative observations are consistent with the hypothesis that New Caledonian crows develop tool‐oriented behaviour because of an increased motivation to perform object combinations that facilitate the necessary learning. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 870–877.     

West Nile Virus and California Breeding Bird Declines

Since it was first detected in 1999, West Nile virus (WNV) quickly spread, becoming the dominant vector-borne disease in North America. Sometimes fatal to humans, WNV is even more widespread among birds, with hundreds of species known to be susceptible to WNV infection in North America alone. However, despite considerable mortality and local declines observed in American crows (Corvus brachyrhynchos), there has been little evidence of a large regional association between WNV susceptibility and population declines of any species. Here we demonstrate a correlation between susceptibility to WNV measured by large-scale testing of dead birds and two indices of overall population change among bird species following the spread of WNV throughout California. This result was due primarily to declines in four species of Corvidae, including all species in this family except common ravens (Corvus corax). Our results support the hypothesis that susceptibility to WNV may have negative population consequences to most corvids on regional levels. They also provide confirmation that dead animal surveillance programs can provide important data indicating populations most likely to suffer detrimental impacts due to WNV.     

Strong between‐site variation in New Caledonian crows’ use of hook‐tool‐making materials

Functional tool use requires the selection of appropriate raw materials. New Caledonian crows Corvus moneduloides are known for their extraordinary tool‐making behaviour, including the crafting of hooked stick tools from branched vegetation. We describe a surprisingly strong between‐site difference in the plant materials used by wild crows to manufacture these tools: crows at one study site use branches of the non‐native shrub Desmanthus virgatus, whereas only approximately 7 km away, birds apparently ignore this material in favour of the terminal twigs of an as‐yet‐unidentified tree species. Although it is likely that differences in local plant communities drive this striking pattern, it remains to be determined how and why crows develop such strong site‐specific preferences for certain raw materials.     

Corvidae are able to understand the logical structure in string-pulling tasks

The ability of the Corvidae to understand the logical structure in string-pulling tasks was studied in a set of experiments with varied position of strings. It was demonstrated that some hooded crows (Corvus cornix L.) and common ravens (Corvus corax L.) successfully completed the tasks where the strings were not intersected but placed so that the bait was positioned opposite the forepart of the empty string. Hooded crows also solved the task where the baits were attached to both strings, but one of the strings was disrupted. The task with two intersected strings where the bait was positioned opposite the forepart of the empty string was not solved by the crows. The results suggest the ability of both examined species to grasp the logical structure of such kind of tasks.     

New data on the brain and cognitive abilities of birds

New evidence of functional analogies and homologies of avian and mammalian brains is presented, as is a revised nomenclature of the most important brain structures. Comparative characteristics of the avian brain and criteria for its progressive development in the phylogeny have been considered. We studied the possibility to use Portmann??s index as one of the indicators of brain development in different avian species. We substantiated the necessity to chose for investigation new sets of avian species with medium (Parus caeruleus and Loxia curvirostra) and low (Larus glaucescens) levels of brain complexity to maintain fully valuable grounds for comparing the cognitive abilities in birds. The main experimentally supported proofs of the existence of elementary thinking and some other cognitive functions in the higher birds have been reviewed. The high levels of cognitive processes that underlie the tool using ability in birds, as well as the similarity to those processes in apes, have been demonstrated from the results obtained in the first decade of the 21st century. Comparative studies on proto-instrumental activity confirmed the ability of hooded crows and ravens to find urgent solution of tool-using tasks. Although birds with a medium level of brain complexity display seemingly rational behavior, it is plausible that they use simpler rules being unable to understand the task logic. It was shown that birds of different orders with a high level of brain complexity demonstrate similar dynamics in the development of abstract concepts. Crossbills, which have a medium level of brain complexity, were able to develop the same concepts at a lower level than the corvids; whereas the seagulls and pigeons, which possess a low level of cognitive abilities, were not able to operate any abstractions and were incapable of solving other cognitive tests. The fact that corvids, parrots, and apes have similar abilities to solve some cognitive tasks supports the hypothesis of the convergent evolution of the brain and cognition in birds and primates.     

Lateralized tool use in wild New Caledonian crows

Many vertebrate species exhibit sensory and motor asymmetries. Laterality studies of tool use have focused on primates, where hemispheric asymmetries, manifested behaviourally in hand preferences, are thought to be associated with complex motor tasks. Here we report strong individual lateralization for tool use in birds. New Caledonian crows, Corvus moneduloides, hold stick tools with their bills while foraging, often with the nonworking end laterally positioned on one side of the head and the working end possibly positioned in the binocular field. We observed four wild crows to determine whether tools were consistently held on one side. All crows showed a significant preference (two right, two left). This preference was independent of any asymmetry in tool manufacture and held for artificial holes similarly accessible for tools held on either side. This is the first demonstration of lateralized tool use in a nonprimate. In addition, all 173 tools used unilaterally were held only on a crow's preferred side. Such pronounced individual laterality for tool use in natural conditions has previously been reported only in humans and chimpanzees.     

Movements of Juvenile Common Ravens in an Arid Landscape

ABSTRACT Movement patterns of juvenile birds are poorly understood, yet critically important ecological phenomena, especially for species with a prolonged juvenile period. We evaluated postfledging movements of juvenile common ravens (Corvus corax) in a western Mojave Desert landscape composed of a mosaic of natural and anthropogenic elements. Generally, ravens do not begin breeding until after their fourth year. We marked 2 annual cohorts of juvenile ravens and followed them from dispersal from their natal territory for up to 33 months. Movements of juvenile common ravens were similar for males and females. Conspecifics and confined livestock feeding operations represented important resources for juvenile ravens, and juveniles were rarely located in open desert. However, initial movements from the natal territory to the nearest communal point subsidy rather than the closest anthropogenic resource suggested juvenile dispersal was influenced by the combination of conspecifics and anthropogenic resources, rather than the distribution of those resources. Land managers concerned with growing raven populations should reduce access to concentrated anthropogenic resources such as landfills and dairies, which serve as important resources for juveniles. Because juvenile ravens rarely venture into open desert, reducing their numbers by lethal removal or other means is unlikely to lessen raven predation of desert tortoises (Gopherus agassizii).     

Ravens judge competitors through experience with play caching

Complex social behavior builds on the mutual judgment of individuals as cooperation partners and competitors [1]. Play can be used for assessing the others' dispositions in humans and nonhuman mammals [2], whereas little is known about birds. Recently, food-caching corvids have been found to rival primates in their ability to judge the behaviors and intentions of others in competition for hidden food [3]. Here, we show that ravens Corvus corax quickly learn to assess the competitive strategies of unfamiliar individuals through interactions with them over caches with inedible items and subsequently apply this knowledge when caching food. We confronted birds with two human experimenters who acted differently when birds cached plastic items: the pilferer stole the cached objects, whereas the onlooker did not. Birds responded to the actions of both experimenters with changing the location of their next object caches, either away from or toward the humans, as if they were testing their pilfering dispositions. In contrast, ravens instantly modified their caching behavior with food, preventing only the competitive human from finding the caches. Playful object caching in a social setting could thus aid ravens in evaluating others' pilfering skills.     

Diversification and biogeographic patterns in four island radiations of passerine birds

Declining diversification rates over time are a well-established evolutionary pattern, often interpreted as indicating initial rapid radiation with filling of ecological niche space. Here, we test the hypothesis that island radiations may show constant net diversification rates over time, due to continued expansion into new niche space in highly dispersive taxa. We investigate diversification patterns of four passerine bird families originating from the Indo-Pacific archipelagos, and link these to biogeographic patterns to provide independent indications of niche filling. We find a declining diversification rate for only one family, the Paradisaeidae (41 species). These are almost completely restricted to New Guinea, and have on average smaller species ranges and higher levels of species richness within grid cells than the other three families. In contrast, we cannot reject constant diversification rates for Campephagidae (93 species), Oriolidae (35 species), and Pachycephalidae (53 species), groups that have independently colonized neighboring archipelagos and continents. We propose that Paradisaeidae have reached the diversity limit imposed by their restricted distribution, whereas high dispersal and colonization success across the geologically dynamic Indo-Pacific archipelagos may have sustained high speciation rates for the other three families. Alternatively, increasing extinction rates may have obscured declining speciation rates in those three phylogenies.     

Stress associated with group living in a long-lived bird

Many long-lived avian species adopt life strategies that involve a gregarious way of life at juvenile and sub-adult stages and territoriality during adulthood. However, the potential associated costs of these life styles, such as stress, are poorly understood. We examined the effects of group living, sex and parasite load on the baseline concentration of faecal stress hormone (corticosterone) metabolites in a wild population of common ravens (Corvus corax). Corticosterone concentrations were significantly higher in non-breeding gregarious ravens than in territorial adults. Among territorial birds, males showed higher stress levels than their mates. Parasite burdens did not affect hormone levels. Our results suggest a key role of the social context in the stress profiles of the two population fractions, and that group living may be more energetically demanding than maintaining a territory. These findings have implications for understanding hormonal mechanisms under different life styles and may inspire further research on the link between hormone levels and selective pressures modulating gregarious and territorial strategies in long-lived birds.     

Management and Conservation Article

Abstract Despite increased knowledge about environmental toxins and changes in lead use (i.e., the mandated use of nonlead paint, gasoline, and shotgun pellets used for hunting waterfowl on federal lands), lead poisoning continues to occur in terrestrial birds. The degree of exposure and its demographic effect, however, continue to be described, emphasizing the growing concern over lead exposure. We examined 302 blood samples from common ravens (Corvus corax) scavenging on hunter-killed large ungulates and their offal piles to determine if lead rifle-bullet residuum was a point source for lead ingestion in ravens. We took blood samples during a 15-month period spanning 2 hunting seasons. Of the ravens tested during the hunting season, 47% exhibited elevated blood lead levels (≥ 10 μg/dL) whereas 2% tested during the nonhunting season exhibited elevated levels. Females had significantly higher blood lead levels than did males. Our results confirm that ravens are ingesting lead during the hunting season and are likely exposed to lead from rifle-shot big-game offal piles.     

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.     

Social learning spreads knowledge about dangerous humans among American crows

Individuals face evolutionary trade-offs between the acquisition of costly but accurate information gained firsthand and the use of inexpensive but possibly less reliable social information. American crows (Corvus brachyrhynchos) use both sources of information to learn the facial features of a dangerous person. We exposed wild crows to a novel 'dangerous face' by wearing a unique mask as we trapped, banded and released 7-15 birds at five study sites near Seattle, WA, USA. An immediate scolding response to the dangerous mask after trapping by previously captured crows demonstrates individual learning, while an immediate response by crows that were not captured probably represents conditioning to the trapping scene by the mob of birds that assembled during the capture. Later recognition of dangerous masks by lone crows that were never captured is consistent with horizontal social learning. Independent scolding by young crows, whose parents had conditioned them to scold the dangerous mask, demonstrates vertical social learning. Crows that directly experienced trapping later discriminated among dangerous and neutral masks more precisely than did crows that learned through social means. Learning enabled scolding to double in frequency and spread at least 1.2 km from the place of origin over a 5 year period at one site.     

Historical biogeography of an Indo‐Pacific passerine bird family (Pachycephalidae): different colonization patterns in the Indonesian and Melanesian archipelagos

Aim We use molecular‐based phylogenetic methods and ancestral area reconstructions to examine the systematic relationships and biogeographical history of the Indo‐Pacific passerine bird family Pachycephalidae (whistlers). Analysed within an explicit spatiotemporal framework, we elucidate distinct patterns of diversification across the Melanesian and Indonesian archipelagos and explore whether these results may be explained by regional palaeogeological events. We further assess the significance of upstream colonization and its role in species accumulation within the region. Location The Indo‐Pacific region, with an emphasis on the archipelagos on either side of the Australo‐Papuan continent. Methods We used three nuclear and two mitochondrial markers to construct a molecular phylogenetic hypothesis of the Pachycephalidae by analysing 35 of the 49 species known to belong to the family. The programs diva and Mr Bayes were used to reconstruct ancestral area relationships and to examine biogeographical relationships across the family, and beast was implemented to assess the timing of dispersal events. Results We constructed a molecular phylogenetic hypothesis for the Pachycephalidae and estimated divergence times and ancestral area relationships. Different colonization patterns are apparent for the Pachycephalidae in the Indonesian and the Melanesian archipelagos. The Indonesian archipelago was colonized numerous times, whereas one or two colonizations of the Melanesian archipelagos account for the entire diversity of that region. After initial colonization of the Melanesian archipelagos some whistler species recolonized Australia and may have commenced a second round of colonization into Melanesia. Main conclusions The contrasting dispersal patterns of whistlers in archipelagos on either side of the Australo‐Papuan continent are congruent with the arrangement and history of islands in each of the regions and demonstrate that knowledge of palaeogeography is important for an understanding of evolutionary patterns in archipelagos. We also highlight that recolonization of continents from islands may be more common than has previously been assumed.     

Recent origin,active speciation and dispersal for the lichen genus Nephroma (Peltigerales) in Macaronesia

Aim We reconstructed the phylogeny of the lichen genus Nephroma (Peltigerales) to assess the relationships of species endemic to Macaronesia. We estimated dates of divergences to test the hypothesis that the species arose in Macaronesia (neo‐endemism) versus the oceanic archipelagos serving as refugia for formerly widespread taxa (palaeo‐endemism). Location Cosmopolitan with a special focus on the archipelagos of the Azores, Madeira and the Canary Islands. Methods DNA sequences were obtained from 18 species for three loci and analysed using maximum parsimony, maximum likelihood and Bayesian inferences. Divergence dates were estimated for the internal transcribed spacer (ITS)‐based phylogeny using a relaxed molecular clock. Reconstruction of the ancestral geographical range was conducted using the Bayesian 50% majority rule consensus tree under a parsimony method. Results The backbone phylogenetic tree was fully supported, with Nephroma plumbeum as sister to all other species. Four strongly supported clades were detected: the Nephroma helveticum, the N. bellum, the N. laevigatum and the N. parile clades. The latter two share a common ancestor and each includes a widespread Holarctic species (N. laevigatum and N. parile, respectively) and all species endemic to Macaronesia. The data suggest a neo‐endemic origin of Macaronesian taxa, a recent range expansion from Macaronesia of both widespread species, a range expansion limited to the Mediteranean Basin and south‐western Europe for another taxon, and a long dispersal event that resulted in a speciation event in the western parts of North America. Main conclusions The Macaronesian endemic species belong to two sister clades and originated from a most recent common ancestor (MRCA) shared with one widely distributed taxon, either N. parile or N. laevigatum. Estimates of the mean divergence dates suggest that the endemics originated in the archipelagos after the rise of the volcanic islands, along with the ancestor to the now widespread species, which probably expanded their range beyond Macaronesia via long‐distance dispersal. This study provides the first phylogenetic evidence of Macaronesian neo‐endemism in lichenized fungi and provides support for the hypothesis that oceanic islands may serve as a source for the colonization of continents. However, further data are needed to properly assess the alternative hypothesis, namely colonization from western North America.     

Breeding system evolution influenced the geographic expansion and diversification of the core Corvoidea (Aves: Passeriformes)

Birds vary greatly in their life‐history strategies, including their breeding systems, which range from brood parasitism to a system with multiple nonbreeding helpers at the nest. By far the most common arrangement, however, is where both parents participate in raising the young. The traits associated with parental care have been suggested to affect dispersal propensity and lineage diversification, but to date tests of this potential relationship at broad temporal and spatial scales have been limited. Here, using data from a globally distributed group of corvoid birds in concordance with state‐dependent speciation and extinction models, we suggest that pair breeding is associated with elevated speciation rates. Estimates of transition between breeding systems imply that cooperative lineages frequently evolve biparental care, whereas pair breeders rarely become cooperative. We further highlight that these groups have differences in their spatial distributions, with pair breeders overrepresented on islands, and cooperative breeders mainly found on continents. Finally, we find that speciation rates appear to be significantly higher on islands compared to continents. These results imply that the transition from cooperative breeding to pair breeding was likely a significant contributing factor facilitating dispersal across tropical archipelagos, and subsequent world‐wide phylogenetic expansion among the core Corvoidea.     

Long-billed curlew nest site selection and success in the Intermountain West

Grassland birds have experienced steeper population declines between 1966 and 2015 than any other bird group on the North American continent, and migratory grassland birds may face threats in all stages of their annual cycle. The grassland-associated long-billed curlew (Numenius americanus) is experiencing population declines in regional and local portions of their North American breeding range. The nesting period is an important portion of the annual cycle when curlews may face demographic rate limitations from a suite of threats including predators and anthropogenic disturbance. We compared nest sites to random sites within breeding territories to examine nest site selection, and modeled correlates of nesting success for 128 curlew nests at 5 Intermountain West sites. Nest sites were 6 times more likely than random sites to be situated adjacent to existing cowpies. Additionally, curlews selected nest sites with shorter vegetation, and less bare ground, grass, and shrub cover than at random sites within their territories. Nest success varied widely among sites and ranged from 12% to 40% in a season with a mean of 27% for all nests during the 2015 and 2016 seasons. Higher nest success probability was associated with higher curlew densities in the area, greater percent cover of conspicuous objects (cowpies, rocks) near the nest, and higher densities of black-billed magpies (Pica hudsonia) and American crows (Corvus brachyrhynchos) at the site. We also found increased probability of nesting success with increased distance from a nest to the nearest potential perch in that territory. Given the central role of working lands to curlews in much of the Intermountain West, understanding limitations to nesting success in these diverse landscapes is necessary to guide adaptive management strategies in increasingly human-modified habitats. We suggest some grazing and irrigation practices already provide suitable nesting conditions for curlews, and others may require only minor temporal shifts to improve compatibility. © 2019 The Wildlife Society.     

Social context hinders humans but not ravens in a short-term memory task

Using resources shared within a social group—either in a cooperative or a competitive way—requires keeping track of own and others’ actions, which, in turn, requires well-developed short-term memory. Although short-term memory has been tested in social mammal species, little is known about this capacity in highly social birds, such as ravens. We compared ravens (Corvus corax) with humans in spatial tasks based on caching, which required short-term memory of one's own and of others’ actions. Human short-term memory has been most extensively tested of all social mammal species, hence providing an informative benchmark for the ravens. A recent study on another corvid species (Corvus corone) suggests their capacity to be similar to the humans’, but short-term memory skills have, to date, not been compared in a social setting. We used spatial setups based on caches of foods or objects, divided into individual and social conditions with two different spatial arrangements of caches (in a row or a 3 × 3 matrix). In each trial, a set of three up to nine caches was presented to an individual that was thereafter allowed to retrieve all items. Humans performed better on average across trials, but their performance dropped, when they had to keep track of partner's actions. This differed in ravens, as keeping track of such actions did not impair their performance. However, both humans and ravens demonstrated more memory-related mistakes in the social than in the individual conditions. Therefore, whereas both the ravens’ and the humans’ memory suffered in the social conditions, the ravens seemed to deal better with the demands of these conditions. The social conditions had a competitive element, and one might speculate that ravens’ memory strategies are more attuned to such situations, in particular in caching contexts, than is the case for humans.     

Global Genetic Differentiation in a Cosmopolitan Pest of Stored Beans: Effects of Geography,Host-Plant Usage and Anthropogenic Factors

Genetic differentiation can be promoted allopatrically by geographic isolation of populations due to limited dispersal ability and diversification over time or sympatrically through, for example, host-race formation. In crop pests, the trading of crops across the world can lead to intermixing of genetically distinct pest populations. However, our understanding of the importance of allopatric and sympatric genetic differentiation in the face of anthropogenic genetic intermixing is limited. Here, we examined global sequence variation in two mitochondrial and one nuclear genes in the seed beetle Callosobruchus maculatus that uses different legumes as hosts. We analyzed 180 samples from 42 populations of this stored bean pest from tropical and subtropical continents and archipelagos: Africa, the Middle East, South and Southeast Asia, Oceania and South America. For the mitochondrial genes, there was weak but significant genetic differentiation across continents/archipelagos. Further, we found pronounced differentiation among subregions within continents/archipelagos both globally and within Africa but not within Asia. We suggest that multiple introductions into Asia and subsequent intermixing within Asia have generated this pattern. The isolation by distance hypothesis was supported globally (with or without continents controlled) but not when host species was restricted to cowpeas Vigna unguiculata, the ancestral host of C. maculatus. We also document significant among-host differentiation both globally and within Asia, but not within Africa. We failed to reject a scenario of a constant population size in the recent past combined with selective neutrality for the mitochondrial genes. We conclude that mitochondrial DNA differentiation is primarily due to geographic isolation within Africa and to multiple invasions by different alleles, followed by host shifts, within Asia. The weak inter-continental differentiation is most likely due to frequent inter-continental gene flow mediated by human crop trade.     

Campylobacter jejuni infection associated with relatively poor condition and low survival in a wild bird

Campylobacter jejuni is the most common foodborne pathogen in industrialized countries. Most human infections come from contaminated poultry, but wild birds are also known to harbor C. jejuni. Wild birds are often described as asymptomatic carriers, but this assumption is based on domestic poultry research. We studied the effects of C. jejuni infection on body condition and survival of adult and nestling American crows Corvus brachyrhynchos in Davis, California. Previous work demonstrated that more than half of the crows in this population are infected with C. jejuni and that at least some of the isolates carried by crows are similar to those found in domestic animals and humans. In this study, we compared the body condition of infected and uninfected individuals at the time of capture among adults (n = 44; 52% infected) and nestlings (n = 97; 77% infected). We subsequently monitored these banded individuals for up to 3 yr and used mark–recapture survival analysis to estimate relationships between infection status and later survival. We found that adult crows infected with C. jejuni were in poor condition relative to uninfected adults: average body mass of infected birds was 12% lower, whereas average body size did not differ between the two groups. Likewise, apparent survival probability was lower for infected adults. In contrast, nestling body condition, fledging success, and survival did not differ by infection status. This is the first study to document adverse effects of C. jejuni infection in a free‐living, wild bird. If these effects are widespread, C. jejuni exposure may be a cause of conservation concern for some species, especially when human activities increase exposure to infections or introduce novel strains to wild bird populations. Our results add to the growing body of work demonstrating hidden long‐term costs of seemingly mild infections in wild populations.