Ravens adjust their antipredatory responses to con‐ and hetero‐specific alarms to the perceived threat

Heterospecific alarm calls are typically found in situations where multiple species have a common predator. In birds, they are particularly common in mixed mixed‐species flocks. In species with highly developed social and cognitive abilities like corvids, there is the potential for differential responses to heterospecific vs. conspecific calls according to the riskiness of the habitat. We tested the responses of free‐ranging ravens (Corvus corax) to conspecific alarm calls and compared them to heterospecific alarm calls of jackdaws (Corvus monedula). We observed the proportion of ravens leaving the feeding site after the con‐ or hetero‐specific playback was presented in a situation of low threat (wild boar—Sus scrofa enclosure) and high threat of predation (wolf—Canis lupus enclosure). We show that ravens responded to conspecific calls more intensively at the wolves than at the wild boar, but the response to conspecific calls was in both enclosures stronger than to the control (great tit—Parus major song). The response to the heterospecific alarm was also stronger in the wolves’ enclosure, but it did not differ from control in the wild boar enclosure. These findings suggest that ravens are aware of the meaning of the jackdaw alarm calls, but they respond to it only in a situation of high predatory threat (wolves are present). In the wild boar enclosure, the ravens probably consider jackdaws warning against some other predator, very probably harmless to ravens. This interpretation requires further testing, as both enclosures differ also in respect to other parameters like food quality and shelter availability.     

Age Differences in the Response of Willow Tits (Parus montanus) to Conspecific Alarm Calls

Predation is an important mortality factor in wintering birds. To counter this, birds produce alarm calls in the presence of predators which serve to warn conspecifics. In social hierarchical bird flocks, adults survive the winter better than juveniles and therefore survival strategies probably vary with social status. This study examined the differential responses to alarm calls by free-living willow tits, Parus montanus, in dominance-structured winter flocks in Finland. To explore the age-dependent differences in response to conspecific alarm calls, a series with three alarm calls was played to focal adults and juveniles while they sat in the middle section of a spruce branch. Immediately after the playback, juvenile willow tits moved more often, flew longer distances and changed branches more often than did adults. Previous mammal studies have shown that juveniles are more likely to flee than adults after hearing conspecific alarm calls. The current study demonstrates that similar age-dependent responses to conspecific alarm calls occur in birds also. These findings reflect an increased vulnerability to predators or lack of experience of young birds.     

THE FORMATION, STRUCTURE AND FUNCTION OF MIXED-SPECIES INSECTIVOROUS BIRD FLOCKS IN WEST AFRICAN SAVANNA WOODLAND

Mixed-species flocks of birds were observed during the wet season (July to September 1975) in savanna woodland in Ghana. Thirty-four flocks contained birds of 56 species in 20 families, including insectivorous, granivorous, and nectarivorous species, using a wide range of foraging methods. Only two species occurred in more than half the flocks. There was no correlation between the number of flocks joined by a species and its abundance in the community. Among insectivores, but not granivores, the species which joined most flocks were those which habitually occurred in the largest single-species groups. All stages of breeding activity were represented by the various members. Some species joined flocks only while these were passing through their territories. Of the two species which were present most frequently, there were no differences between mixed and single-species flocks for Eremomela pusilla, but Parus leucomelas foraged and called on more occasions in mixed flocks than single-species flocks, though the rates of foraging and calling were related only to the number of P. leucomelas present. Groups of P. leucomelas appeared to initiate some flocks by attraction due to their conspicuous wing-bars, active movement, and loud calls. Black-and-white species joined them first, followed by birds of other plumage patterns. The advantages of mixed flocking are thought to be connected with finding patches of the food of bark- and foliage-searching insectivores, which were the only species regularly seen foraging in the flocks. Because of dry season burning which leaves small unburnt patches of savanna, these insect species may share a common, patchy distribution. Birds may also gain protection from predators, and some species probably gain no advantages. The species composition and behaviour of flocks previously recorded elsewhere in African savannas are similar to the Ghana flocks.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Functional traits determine heterospecific use of risk‐related social information in forest birds of tropical South‐East Asia

In birds and mammals, mobbing calls constitute an important form of social information that can attract numerous sympatric species to localized mobbing aggregations. While such a response is thought to reduce the future predation risk for responding species, there is surprisingly little empirical evidence to support this hypothesis. One way to test the link between predation risk reduction and mobbing attraction involves testing the relationship between species’ attraction to mobbing calls and the functional traits that define their vulnerability to predation risk. Two important traits known to influence prey vulnerability include relative prey‐to‐predator body size ratio and the overlap in space use between predator and prey; in combination, these measures strongly influence prey accessibility, and therefore their vulnerability, to predators. Here, we combine community surveys with behavioral experiments of a diverse bird assemblage in the lowland rainforest of Sumatra to test whether the functional traits of body mass (representing body size) and foraging height (representing space use) can predict species’ attraction to heterospecific mobbing calls. At four forest sites along a gradient of forest degradation, we characterized the resident bird communities using point count and mist‐netting surveys, and determined the species groups attracted to standardized playbacks of mobbing calls produced by five resident bird species of roughly similar body size and foraging height. We found that (1) a large, diverse subcommunity of bird species was attracted to the mobbing calls and (2) responding species (especially the most vigorous respondents) tended to be (a) small (b) mid‐storey foragers (c) with similar trait values as the species producing the mobbing calls. Our findings from the relatively lesser known bird assemblages of tropical Asia add to the growing evidence for the ubiquity of heterospecific information networks in animal communities, and provide empirical support for the long‐standing hypothesis that predation risk reduction is a major benefit of mobbing information networks.     

Bird mixed‐species flock formation is driven by low temperatures between and within seasons in a Subtropical Andean‐foothill forest

According to both the predation avoidance and foraging efficiency hypotheses, birds within mixed flocks increase their foraging efficiency and/or can spend more time feeding and less time looking out for predators. These hypotheses predict that birds in mixed flocks obtain benefits. Thus, mixed flock formation could serve as a strategy to cope with difficult conditions imposed on birds such as climatic conditions that ultimately result in a change in predation pressure or food resources. We evaluate the hypotheses that forming part of a flock confers benefits to its members and the associated prediction that birds will take advantage of these benefits and flock more often under cold and dry weather conditions between and within seasons to cope with such conditions. We surveyed the presence of mixed flocks, flocking propensity, number of species and individuals in mixed flocks in the Subtropical Yungas foothill of Argentina, to examine seasonality, flocking behavior of birds and their responses to two climatic variables: temperature and humidity. Bird species presented a higher flocking propensity and mixed flocks occurred more frequently during the dry and cold seasons than during the more benign seasons, and lower values of temperature within seasons triggered the flocking behavior. Although effects between seasons were expected, birds also showed a short‐term response to small changes in temperature within seasons. These results strengthen the ideas proposed by the foraging hypothesis. Although benefits derived from flocking have yet to be determined, whatever they are should be understood in the context of seasonal variation in life‐history traits.     

Differential Response to Interspecific and Intraspecific Signals Amongst Chickadees

Black‐capped chickadees (Poecile atricapillus) and mountain chickadees (P. gambeli) have a similar vocal repertoire and share many other life history traits; yet, black‐capped chickadees are socially dominant to mountain chickadees where populations overlap. Previous research suggested that in contact zones, both species respond weakly to heterospecific songs during the breeding season, and have suggested minimal interspecific competition. However, both black‐capped and mountain chickadees discriminate between conspecific and heterospecific chick‐a‐dee calls, suggesting attention is paid to interspecific signals. We compared the responses of both black‐capped and mountain chickadees to conspecific and heterospecific chick‐a‐dee calls during the winter, when both species compete for the same food resources. We conducted an aviary playback experiment exposing both species to playback composed of heterospecific and conspecific chick‐a‐dee calls, which had been recorded in the context of finding food sources. Responses from the tested birds were measured by recording vocalizations and behaviour. Black‐capped chickadees responded significantly more to conspecific than to heterospecific stimuli, whereas the subordinate mountain chickadees responded to both mountain and black‐capped chickadee calls. Based upon the reactions to playbacks, our results suggest these two closely related species may differ in their perception of the relative threat associated with intra‐ versus interspecific competitors.     

Behavioral responses to conspecific mobbing calls are predator‐specific in great tits (Parus major)

When facing a predator, animals need to perform an appropriate antipredator behavior such as escaping or mobbing to prevent predation. Many bird species exhibit distinct mobbing behaviors and vocalizations once a predator has been detected. In some species, mobbing calls transmit information about predator type, size, and threat, which can be assessed by conspecifics. We recently found that great tits (Parus major) produce longer D calls with more elements and longer intervals between elements when confronted with a sparrowhawk, a high‐threat predator, in comparison to calls produced in front of a less‐threatening tawny owl. In the present study, we conducted a playback experiment to investigate if these differences in mobbing calls elicit different behavioral responses in adult great tits. We found tits to have a longer latency time and to keep a greater distance to the speaker when sparrowhawk mobbing calls were broadcast. This suggests that tits are capable of decoding information about predator threat in conspecific mobbing calls. We further found a tendency for males to approach faster and closer than females, which indicates that males are willing to take higher risks in a mobbing context than females.     

Eavesdropping on heterospecific alarm calls: from mechanisms to consequences

Animals often gather information from other species by eavesdropping on signals intended for others. We review the extent, benefits, mechanisms, and ecological and evolutionary consequences of eavesdropping on other species' alarm calls. Eavesdropping has been shown experimentally in about 70 vertebrate species, and can entail closely or distantly related species. The benefits of eavesdropping include prompting immediate anti‐predator responses, indirect enhancement of foraging or changed habitat use, and learning about predators. Eavesdropping on heterospecifics can provide more eyes looking for danger, complementary information to that from conspecifics, and potentially information at reduced cost. The response to heterospecific calls can be unlearned or learned. Unlearned responses occur when heterospecific calls have acoustic features similar to that used to recognize conspecific calls, or acoustic properties such as harsh sounds that prompt attention and may allow recognition or facilitate learning. Learning to recognize heterospecific alarm calls is probably essential to allow recognition of the diversity of alarm calls, but the evidence is largely indirect. The value of eavesdropping on different species is affected by problems of signal interception and the relevance of heterospecific alarm calls to the listener. These constraints on eavesdropping will affect how information flows among species and thus affect community function. Some species are ‘keystone’ information producers, while others largely seek information, and these differences probably affect the formation and function of mixed‐species groups. Eavesdroppers might also integrate alarm calls from multiple species to extract relevant and reliable information. Eavesdropping appears to set the stage for the evolution of interspecific deception and communication, and potentially affects communication within species. Overall, we now know that eavesdropping on heterospecific alarm calls is an important source of information for many species across the globe, and there are ample opportunities for research on mechanisms, fitness consequences and implications for community function and signalling evolution.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

The Payoffs to Producing and Scrounging: What Happens when Patches are Divisible?

Although many group-foraging models assume that all individuals search for and share their food equally, most documented instances of group foraging exhibit specialized use of producer and scrounger strategies. In addition, many of the studies have focused on groups with strong individual asymmetries exploiting food that is not easily divisible. In the present study we describe individual foraging behavior of relatively nonaggressive flock foragers exploiting divisible clumps of food. Two experiments, one with flocks of spice finches and another with flocks of zebra finches, suggest that divisibility of food patches may have important consequences for social foraging behavior. Neither dominance nor the distribution and quality of food patches affect the relative advantage that producing individuals enjoy over those that scrounge. Specialized producers and scroungers are absent from flocks of both species. Systems where patches are shared may differ fundamentally from those where patches are monopolized by scroungers.     

Numerical and behavioural responses of migrant passerines to experimental manipulation of resident tits (Parus spp.): heterospecific attraction in northern breeding bird communites?

Summary We studied experimentally interspecific competition among foliage-gleaning passerine birds by manipulating the density of resident tits. In 1988 tit density was experimentally increased on three small islands in a central Finnish lake, and decreased on three other islands by tit removal. In order to avoid the effects of between-island differences in habitat quality, the role of the islands was reversed when the experiment was repeated in the following year. Censuses and observations on foraging and feeding behaviour were conducted to assess the numerical and behavioural responses of migrant conguilders (mainly chaffinches and willow warblers) with respect to the manipulated abundance of the tits. We also measured whether variation in food consumption of tits affected the frequency with which the migrants found food by calculating average intervals between successful prey captures, time lags to prey-capture and giving-up times. Our results indicate that interspecific competition is of minor importance in structuring breeding bird assemblages and species feeding ecologies on the study islands. No consistent difference in foraging or feeding niches of chaffinches and willow warblers was found between low and high tit density conditions. Niche overlap analysis showed no avoidance by chaffinches and willow warblers of the microhabitats which tits used. Tit abundance had no significant effect on feeding success or behaviour. Experimentally increased abundance of resident birds was associated with increased abundance of breeding migrants, however. This pattern was found not only in the foliage gleaning guild but also with all passerine birds, indicating that food was not an important contributor to this pattern. We elaborate a hypothesis suggesting heterospecific attraction in northern breeding bird assemblages. Habitat generalist migrants may use the presence of residents as an indicator of safe and/or productive breeding sites in northern unpredictable circumstances.     

Alarm calls of wintering great tits Parus major: warning of mate,reciprocal altruism or a message to the predator?

When a predator is not an immediate threat, a prey may produce relatively loud alarm calls because the risk is low. Since such calls could nevertheless attract acoustically oriented predators, the cost of predator attraction must be outweighed by factors beneficial to the caller. In this field study we elicited low-risk alarm calls by temporarily catching wintering adult male great tits Parus major at feeders both within and outside their territories. We tested whether the alarm calls of dominant males can be explained in terms of mate warning, reciprocal altruism or notifying the predator of detection. If alarms are intended to warn mates, males accompanied by their mates should give alarm calls both within and outside home range, even if other permanent flock members are absent. If alarms are to be explained by reciprocal altruism, male great tits should give low-risk alarm calls when accompanied by permanent flock members other than mate within and not outside of the home-range. If alarm calling is a message to a predator, males should call when foraging alone. We found that male great tits gave low-risk alarm calls when accompanied by their mates, independent of feeder location. They also gave low-risk alarm calls within home ranges in the presence of other permanent flock members when mates were absent. In contrast, only a few males gave calls when foraging alone within their home ranges, or when in the company of unfamiliar great tits outside their usual home-range. The results suggest that the utterance of alarm calls may be explained as mate protection and reciprocal altruism among familiar individuals.     

False Alarm Calls as a Means of Resource Usurpation in the Great Tit Parus major

Great tits Parus major regularly gave alarm calls in winter without the presence of actual or potential predators. Such false alarm calls were given to deceive both conspecifics and heterospecifics. False calls were given when flock-feeding sparrows Passer spp. monopolized a concentrated food source; if the food resource was dispersed false alarm calls were not used, independently of the presence of sparrows. Dominant great tits used alarm calls deceptively if a dominant conspecific was present on a concentrated food source but not if a subdominant individual was present; subdominant great tits were displaced by means of threat displays. Subdominant individuals gave false alarm calls both if dominant or subdominant conspecifics were present. False alarm calls were especially used when food was scarce or when great tits were feeding at a high rate, i.e. during snow storms and in the morning and the afternoon.     

Observational and Experimental Evidence for the Function of Tail Flicking in Eurasian Moorhen Gallinula chloropus

Tail movements such as wagging, flicking or pumping are reported from many bird species but their adaptive functions remain poorly understood. To investigate whether tail flicking functions as an alarm signal, either to predators or neighbouring birds, or as a signal of submission to conspecifics, I observed this behaviour in moorhen in a natural context, and conducted playback experiments using vocalizations of predators, conspecifics and heterospecifics. I found positive relationships between flicking and vigilance and nearest neighbour distance, and negative relationships between flicking and moorhen flock size and total flock size. Moorhen at the edge of a flock flicked at a higher rate. Single moorhen flicked more often compared with individuals in groups, both in single‐ and mixed‐species flocks, and there was a tendency that single moorhen flicked more often than single moorhen within a mixed‐species flock. Moorhen responded differently to conspecific and predator calls. While in both cases vigilance increased, tail flick rate was higher during predator playbacks and lower during conspecific playbacks. Furthermore, moorhen remained rather motionless when conspecific calls were played back, but not during predator calls, and, moorhen resumed to a baseline level of tail flicking more quickly after the playback of conspecific calls. Taken together, the results suggest that flicking may be considered as a honest signal of vigilance directed towards ambushing predators.     

Simulated chorus attracts conspecific and heterospecific Amazonian explosive breeding frogs

In tropical regions, some anuran species breed "explosively", reproducing in massive and highly diverse aggregations during a brief window of time. These aggregations can serve as acoustic beacons, attracting other anurans toward seasonal ponds. We hypothesize that conspecific and heterospecific calls play a role in navigation toward ponds and synchronization of reproduction among species. We simulated a chorus of two species (Trachycephalus coriaceus and Chiasmocleis shudikarensis) with contrasting call characteristics (low‐frequency vs. high‐frequency) and reproductive strategies (strict pond breeder vs. opportunistic) near known explosive breeding sites. We predicted that choruses of T. coriaceus are more attractive to heterospecifics than of C. shudikarensis because the first provides a more reliable indicator of a suitable breeding pond and a better long‐distance signal. We found that both choruses attracted conspecific frogs to the playback outside a natural breeding event. As predicted, heterospecifics were attracted only by low‐frequency calls of T. coriaceus that breeds exclusively in large ponds, but not by higher frequency calls of C. shudikarensis that also breeds in small pools not suitable for other species. Our study presents the first experimental evidence that tropical explosive breeding anurans are attracted to conspecific and heterospecific choruses. The contrasting effect of the playback of the two species on heterospecifics suggests that the attractive effect of a chorus depends on the reproductive strategy of both the sender and the receiver. Given the abundance and diversity of communities in tropical ecosystems, the use of heterospecific acoustic cues may prove widespread and requires further investigation. Abstract in Portuguese is available with online material.     

Mycorrhizal feedback is not associated with the outcome of competition in old‐field perennial plants

The symbiosis between plants and arbuscular mycorrhizal (AM) fungi is hypothesized to be an important contributor to plant–soil feedbacks, which can influence the outcome of inter‐specific competition. Mycorrhizal feedbacks can be conspecific, which affects individuals of the same species, or heterospecific, which affects individuals of a different species. When heterospecific feedbacks are more positive than conspecific feedbacks, heterospecific individuals are expected to outcompete conspecific individuals. To test this hypothesis, we quantified conspecific mycorrhizal feedback for Plantago lanceolata as a focal species, and heterospecific mycorrhizal feedbacks for 21 competitor old‐field species using mycorrhizae cultured with P. lanceolata. We quantified inter‐specific competition against the focal species by growing the 21 old‐field species with and without P. lanceolata in the presence of mycorrhizae cultured with P. lanceolata. Heterospecific and conspecific feedbacks were both positive, and average heterospecific feedbacks exceeded conspecific feedback by 75%. Competition suppressed P. lanceolata biomass by 14% and average competitor biomass was reduced by 44% in the presence of P. lanceolata, and these effects varied with competitor species identity. Contrary to predictions, the magnitude of heterospecific feedbacks did not predict the ability of competitor species to either suppress or resist suppression by P. lanceolata. Instead, the outcome of competition was significantly and positively correlated with intrinsic growth rate, measured as biomass of competitor species five weeks after germination in non‐inoculated conditions. Our findings suggest that species experiencing more positive mycorrhizal feedbacks than a competitor do not necessarily have a competitive advantage. Mycorrhizal mediated soil feedbacks may be less important than intrinsic differences in growth rate in determining competitive outcomes.     

Alarm call‐based discrimination between common cuckoo and Eurasian sparrowhawk in a Chinese population of great tits

Morphological resemblance of the common cuckoo Cuculus canorus to the Eurasian sparrowhawk Accipiter nisus has been regarded as an example of predator mimicry. Common hosts could distinguish parasites as the result of coevolution, while rare hosts or non‐hosts may mistake cuckoos for hawks because rare hosts or non‐hosts behave similarly when faced with these two species. Birds usually produce alarm calls in addition to showing behavioral responses when in danger. However, previous studies of identification by rare hosts or non‐hosts of sparrowhawks usually lacked experimental evidence of alarm calls. Great tits Parus major, a rare cuckoo host, perform similar behaviors and usually produce alarm calls in response to sparrowhawks and common cuckoos. Here, we tested whether great tits could distinguish common cuckoo from sparrowhawk based on analysis of their alarm calls and the effects of playback of alarm calls on conspecific behavior. Previous studies showed that great tits have a complex communication system that conveys information about predators, and they could perform different kinds of response behavior to different alarm calls. If great tits have not made the ability to distinguish between common cuckoo and sparrowhawk, then their acoustic responses to these two species and their response behaviors in playback experiments should be similar. Specimens of a common cuckoo (parasite), a sparrowhawk (predator) and an Oriental turtle dove Streptopelia orientalis (harmless control) were used to elicit and subsequently record the response behavior and alarm calls of great tits. There was no significant difference in behavioral response among great tits when exposed to the dummy of cuckoo, sparrowhawk and dove. In contrast, they differed significantly in alarm calls. Great tits produced more notes per call that contained increasing D‐type and decreasing I‐type notes when responding to sparrowhawk as compared to cuckoo or dove. In playback experiments, we found that great tits responded more strongly to great tit hawk than to great tit cuckoo or great tit dove alarm calls. Our study suggests that great tits are able to distinguish sparrowhawks from common cuckoos and convey relevant information in alarm calls by adjusting the number and combinations of notes of a single call type.     

Does predation maintain tit community diversity?

European tits of the genus Parus constitute a complex group of coexisting boreal birds. Here we present a survey of the distribution of three coniferous-living Parus species and one of their main predators, the pygmy owl ( Glaucidium passerinum ), on nine isolated islands in Scandinavia. On all islands the coal tit ( Parus ater ) is the sole tit species when the pygmy owl is absent. The two larger species, the willow tit ( P. montanus ) and the crested tit ( P. cristatus ), only coexist with the coal tit when pygmy owls are present. We suggest that the coexistence of willow tits, crested tits and coal tits is the result of a combination of competition for food and predator-safe foraging sites. The smaller coal tit is superior in exploitation competition for food, while the two larger species have an advantage in interference competition for predator-safe foraging sites. The association between the distribution of the pygmy owl and the two larger tit species on isolated islands in Scandinavia is consistent with the idea that the pygmy owl is a keystone predator.     

Asymmetrical interspecific communication of predatory threat in mixed‐species colonies of lesser kestrels (Falco naumanni) and jackdaws (Corvus monedula)

Sympatric species derive benefits by attending to information conveyed by heterospecifics. Our previous finding of reduced vigilance among jackdaws and lesser kestrels residing in mixed‐species colonies suggested a reliance on interspecific communication of information regarding predatory threats. To test for interspecific communication of threat, we first determined whether jackdaw and lesser kestrel call structure varied with perceived threat. In this call production phase of our study, free‐living birds in mixed‐species colonies were presented with models representing a potential nest predator (European magpie) or with non‐threatening stimuli (wood pigeon or wooden dowel) in proximity to nests. We recorded and subsequently analysed those calls to determine if any temporal or frequency‐related call parameters differed by model type. In a second, perceptual phase of our study, we tested whether receivers perceive threat‐related variation in both conspecific and heterospecific call structure by playing back call exemplars recorded in response to the predator model or to innocuous control stimuli, to determine whether free‐living jackdaws or lesser kestrels respond differentially to playbacks of the different call types. We detected differences in vocalizations of both jackdaws and lesser kestrels relative to the model type presented, with more broadband (lesser kestrel) or noisy calls (jackdaws) in response to magpie versus innocuous model types. We also detected differential behavioural responses to call playbacks, with both jackdaws and lesser kestrels increasing vigilance and alarm calling in response to magpie‐elicited jackdaw calls, but not to other call types. Taken together, our results suggest that jackdaw, but not lesser kestrel vocalizations, communicate enhanced threat associated with European magpies as possible nest predators. This interspecific alarm communication benefits both jackdaws and lesser kestrels, and, at least in part, explains asymmetric responses of jackdaws and lesser kestrels to magpies attending mixed‐species colonies in nature.