Increased Noise Levels Have Different Impacts on the Anti-Predator Behaviour of Two Sympatric Fish Species

Animals must avoid predation to survive and reproduce, and there is increasing evidence that man-made (anthropogenic) factors can influence predator−prey relationships. Anthropogenic noise has been shown to have a variety of effects on many species, but work investigating the impact on anti-predator behaviour is rare. In this laboratory study, we examined how additional noise (playback of field recordings of a ship passing through a harbour), compared with control conditions (playback of recordings from the same harbours without ship noise), affected responses to a visual predatory stimulus. We compared the anti-predator behaviour of two sympatric fish species, the three-spined stickleback (Gasterosteus aculeatus) and the European minnow (Phoxinus phoxinus), which share similar feeding and predator ecologies, but differ in their body armour. Effects of additional-noise playbacks differed between species: sticklebacks responded significantly more quickly to the visual predatory stimulus during additional-noise playbacks than during control conditions, while minnows exhibited no significant change in their response latency. Our results suggest that elevated noise levels have the potential to affect anti-predator behaviour of different species in different ways. Future field-based experiments are needed to confirm whether this effect and the interspecific difference exist in relation to real-world noise sources, and to determine survival and population consequences.     

Responses of Kamchatkan fish‐eating killer whales to playbacks of conspecific calls

Killer whales produce repertoires of stereotyped call types that are primarily transmitted vertically through social learning, leading to dialects between sympatric pods. The potential function of these call repertoires remains untested. In this study, we compared the reaction of Kamchatkan fish‐eating killer whales to the playbacks of calls from the same and different pods. After the playback of recordings from a different pod, in three cases whales changed the direction of their movement toward the boat, and in three cases no changes in direction were observed. After the playback of recordings from the same pod (either from the same or a different unit within the pod), in seven cases whales changed the direction of their movement toward the boat, and in only one case no change in direction was observed. Whales remained silent after all six playbacks of recordings from a different pod, even when they changed direction toward the boat. After the playback of recordings from the same pod, however, in all eight cases whales started calling in response. Our playback study shows that killer whales may react to playbacks of conspecific sounds and that reactions are dependent on the type of playback stimuli.     

The effects of anthropogenic sources of sound on fishes

There is increasing concern about the effects of pile driving and other anthropogenic (human-generated) sound on fishes. Although there is a growing body of reports examining this issue, little of the work is found in the peer-reviewed literature. This review critically examines both the peer-reviewed and 'grey' literature, with the goal of determining what is known and not known about effects on fish. A companion piece provides an analysis of the available data and applies it to estimate noise exposure criteria for pile driving and other impulsive sounds. The critical literature review concludes that very little is known about effects of pile driving and other anthropogenic sounds on fishes, and that it is not yet possible to extrapolate from one experiment to other signal parameters of the same sound, to other types of sounds, to other effects, or to other species.     

A field test of the audibility of urban versus rural songs in mountain chickadees

Anthropogenic noise can mask avian vocalizations, and several urban‐dwelling species adjust frequency or amplitude of vocalizations in ways that appear to compensate for increased noise levels. Playback studies have investigated whether receivers differentiate between signals produced by rural and urban males, but it is difficult to determine whether differential response to stimulus reflects differences in audibility versus perceived differences in male signals/condition that result from urban settlement. Here, we performed paired‐playback trials to determine whether mountain chickadees (Poecile gambeli) differentiate urban versus rural songs when both stimuli were broadcast within noise. For each playback, stimuli were played in short bouts starting at a low signal‐to‐noise ratio and increasing in relative amplitude with each successive bout. If the primary function of adjusted urban songs is propagation in noise, we hypothesized that focal males would respond sooner to urban versus rural playbacks (detect at lower signal‐to‐noise ratios). If urban songs solely encode information about the male's condition (such as increased aggression), then we predicted only a differential aggressive response to playbacks once detected. If urban songs both increase propagation and embed information on male condition, we predicted a combination of both response types. We found no difference in latency to first response in urban versus rural songs, but some evidence for differential aggression to playback dependent on both stimulus type (urban vs. rural) and local ambient noise levels; focal males in noisy (urban) sites responded aggressively to both stimulus types, whereas focal males in quiet (rural) sites responded more aggressively to urban than to rural stimuli. This context‐dependent discrimination may be the result of increased aggression in urban habitats, improved communication in noisy habitats by urban signallers and receivers, or some combination of the two.     

Little Evidence to Support the Risk–Disturbance Hypothesis as an Explanation for Responses to Anthropogenic Noise by Pygmy Marmosets (Cebuella niveiventris) at a Tourism site in the Peruvian Amazon

The risk–disturbance hypothesis states that animals react to human stressors in the same way as they do to natural predators. Given increasing human–wildlife contact, understanding whether animals perceive anthropogenic sounds as a threat is important for assessing the long-term sustainability of wildlife tourism and proposing appropriate mitigation strategies. A study of pygmy marmoset (Cebuella niveiventris) responses to human speech found marmosets fled, decreased feeding and resting, and increased alert behaviors in response to human speech. Following this study, we investigated pygmy marmoset reactions to playbacks of different acoustic stimuli: controls (no playback, white noise and cicadas), anthropogenic noise (human speech and motorboats), and avian predators. For each playback condition, we recorded the behavior of a marmoset and looked at how the behaviors changed during and after the playback relative to behaviors before. We repeated this on ten different marmoset groups, playing each condition once to each group. The results did not replicate a previous study on the same species, at the same site, demonstrating the importance of replication in primate research, particularly when results are used to inform conservation policy. The results showed increased scanning during playbacks of the cicadas and predators compared with before the playback, and an increase in resting after playbacks of avian predators, but no evidence of behavior change in response to playbacks of human speech. There was no effect of ambient sound levels or distance between the playback source and focal animals on their behavior for all playback conditions. Although we find that noise can change the behavior of pygmy marmosets, we did not find evidence to support the risk–disturbance hypothesis.

     

Dual Function of Chip Calls Depending on Changing Call Rate Related to Risk Level in Territorial Pairs of White‐Eared Ground‐Sparrows

Different mechanisms have been proposed for encoding information into vocalizations: variation of frequency or temporal characteristics, variation in the rate of vocalization production, and use of different vocalization types. We analyze the effect of rate variation on the dual function of chip calls (contact and alarm) produced by White‐eared Ground‐sparrows (Melozone leucotis). We conducted an acoustic playback experiment where we played back 1 min of four chip call rates (12, 36, 60, 84 calls/min). We measured the response of territorial pairs using behavioral responses, and fine structural features of calls produced in response to those playbacks. White‐eared Ground‐sparrows showed more intense behavioral responses to higher than lower call rate playbacks. Both individuals of the pair approached the source of the playback stimulus faster, produced the first vocalization faster, produced more vocalizations, and spent more time close to the stimulus in higher call rate than in lower call rate playbacks. Frequency and duration characteristics of calls (chip and tseet) were similar in response to all call rate playbacks. Our playback experiment elicited different intensity of behavioral responses, suggesting that risk‐based information is encoded in call rate. Our results suggest that variation in the rate of chip call production serves a dual function in this species; calls are used at lower rates for pair contact and at higher rates for alarm/mobbing signals.     

Mating interference of glassy‐winged sharpshooters,Homalodisca vitripennis

Animal communication is a complex behavior that is influenced by abiotic and biotic factors of the environment. Glassy‐winged sharpshooters (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), primarily use vibrational signaling for courtship communication. Because GWSS is a major pest, transmitting the plant pathogenic bacterium Xylella fastidiosa Wells et al., interruption of communication is a possible avenue for control. Playback of white noise, pre‐recorded female signals, and artificial female noise (continuously overlapping female signals) significantly reduced mating of GWSS when compared to silent control mating trials. Furthermore, to begin to determine the mechanism underlying playback control, female signaling activity was recorded in the presence of stimuli. In response to playback of female signals, females signaled (duet‐like) more often than females tested in the absence of playback. After the first playback, almost two‐thirds of females signaled a response within 3 s. Additionally, one‐third of the females signaled within 1 s after cessation of white noise, and significantly more in the time periods following noise termination. Results highlight how GWSS responds to differing competitive disturbances in the environment and lays important ground work that possibly could be used to develop pesticide‐free control methods.     

Shift down,look up: A test of the non‐linearity and fear hypothesis in a non‐vocal skink

The non‐linearity and fear hypothesis predicts that certain non‐linear sounds are one way to evoke antipredator responses in both birds and mammals. This hypothesis, however, has not been studied in non‐vocal species or in reptiles. Such a study would be important because if non‐linear sounds are evocative even in a species that does not produce sounds, then there may be generally salient cues of risk in these sounds. We asked whether non‐vocal lizards, white‐bellied copper‐striped skinks (Emoia cyanura), respond to experimentally broadcast non‐linearities. This species is ideal to ask the question in because prior research has shown that they respond to predator sounds and alarm calls of other species even though they are not vocal. We conducted playback experiments with three computer‐generated simulated non‐linearities to assess whether or not skinks increased antipredator behavior after hearing them. We controlled for novelty by broadcasting a 3‐kHz, 500‐ms pure tone and tropical kingbird (Tyrannus melancholicus) song. Our treatments consisted of a 3‐kHz, 400‐ms pure tone followed by a frequency shift up to 5‐kHz for 100‐ms, a 3‐kHz, 400‐ms pure tone to frequency shift down to 1‐kHz for 100‐ms, and a pure tone followed by 100‐ms of white noise. Following a total of 222 playbacks, we categorized responses into looking, locomotion, and high locomotion, focusing on how skinks changed their rates of time allocation from baseline. We examined 95% confidence intervals to identify whether skinks responded to playbacks and fitted general linear models followed by pairwise comparisons to ask whether skinks discriminated between broadcast stimuli. We found that skinks were especially responsive to frequency downshifts: They significantly increased looking and locomotion, consistent with our predictions based on the non‐linearity and fear hypothesis. Surprisingly, they decreased rates of looking behavior after hearing frequency upshifts, possibly suggesting an increase in relaxed behavior. While skinks responded to noise by increasing their rate of locomotion, this response was not significantly different from controls. We conclude that skinks increase antipredator behavior after hearing downshifts more than any other type of non‐linearity. This provides some support for the non‐linearity and fear hypothesis; even non‐vocal species may respond fearfully to specific types of non‐linear sounds.     

Complex Vocal Responses to Conspecific Calls in Whistling Frogs Eleutherodactylus johnstonei: Playback Experiments in the Field

We simulated the presence of an acoustic competitor by broadcasting conspecific playbacks to males of Johnstone's whistling frog, Eleutherodactylus johnstonei, in the field. We broadcast calls that differed in duration (short, typical, and long), dominant frequency (high, typical, and low), and period (short, typical, and long), and analyzed male vocal responses. We tested the hypothesis that males respond by escalating vocally when they are exposed to female‐attractive calls and by ignoring unattractive ones. At the population level, males responded to playbacks in ways that would potentially increase their attractiveness with regard to solo calling: males increased the duration, reduced the dominant frequency, and increased their calling effort (duty cycle), despite an increase in call period. The modification of call duration occurred only in response to playbacks of low‐frequency calls, long calls, and short‐period calls (selective response), while the modification of the dominant frequency was independent of the characteristic of the playback (fixed response). Contrary to the expected, males did not reduce the call period when they were exposed to attractive playbacks. At the ultimate level, the results suggest energy‐saving strategies. In addition, males seem to trade off call period for the avoidance of acoustic interference with attractive calls as calling effort was typically increased by increasing call duration but only rarely by reducing the call period. Interactive playbacks are necessary to better understand the calling strategies of males of E. johnstonei.     

A simulation approach to assessing environmental risk of sound exposure to marine mammals

Intense underwater sounds caused by military sonar, seismic surveys, and pile driving can harm acoustically sensitive marine mammals. Many jurisdictions require such activities to undergo marine mammal impact assessments to guide mitigation. However, the ability to assess impacts in a rigorous, quantitative way is hindered by large knowledge gaps concerning hearing ability, sensitivity, and behavioral responses to noise exposure. We describe a simulation‐based framework, called SAFESIMM (Statistical Algorithms For Estimating the Sonar Influence on Marine Megafauna), that can be used to calculate the numbers of agents (animals) likely to be affected by intense underwater sounds. We illustrate the simulation framework using two species that are likely to be affected by marine renewable energy developments in UK waters: gray seal (Halichoerus grypus) and harbor porpoise (Phocoena phocoena). We investigate three sources of uncertainty: How sound energy is perceived by agents with differing hearing abilities; how agents move in response to noise (i.e., the strength and directionality of their evasive movements); and the way in which these responses may interact with longer term constraints on agent movement. The estimate of received sound exposure level (SEL) is influenced most strongly by the weighting function used to account for the specie's presumed hearing ability. Strongly directional movement away from the sound source can cause modest reductions (~5 dB) in SEL over the short term (periods of less than 10 days). Beyond 10 days, the way in which agents respond to noise exposure has little or no effect on SEL, unless their movements are constrained by natural boundaries. Most experimental studies of noise impacts have been short‐term. However, data are needed on long‐term effects because uncertainty about predicted SELs accumulates over time. Synthesis and applications. Simulation frameworks offer a powerful way to explore, understand, and estimate effects of cumulative sound exposure on marine mammals and to quantify associated levels of uncertainty. However, they can often require subjective decisions that have important consequences for management recommendations, and the basis for these decisions must be clearly described.     

Effects of experimental playbacks on availability for detection during point counts

Point counts are the most commonly used technique for surveying passerines during the breeding season. Several methods for estimating probabilities of detection during point count surveys have been developed. These methods have focused primarily on accounting for the influence of environmental factors (e.g., weather and noise) on detectability, however, the probability that birds are available for detection (e.g., sings or moves) during point counts has received less attention. We used sequential point counts to determine the effect of playback of the mobbing calls of Black‐capped Chickadees (Poecile atricapillus) and the flight calls of Red‐tailed Hawks (Buteo jamaicensis) on availability for detection (e.g., singing or moving) during point‐count surveys. We conducted 180 point counts over a 2‐yr period in central – east central Minnesota to evaluate the possible effect of playbacks on observed density, overall species richness, minute of first detection, and distance of first detection. We also used removal models to quantify the magnitude of changes in detectability and direction of response to playbacks for 10 focal species. Playback of the mobbing calls of Black‐capped Chickadees increased observed density and decreased the average distance of detection and time of first detection, whereas playback of the flight calls of a Red‐tailed Hawk resulted in a decrease in observed density and species richness, and an increased time of first detection. Playback treatment was a covariate in all best performing models for the 10 species analyzed, but the magnitude and direction of response to playbacks were species specific. The importance of playback type in detectability models indicates that the calls of heterospecifics can influence species availability for detection. As such, researchers using playback methods should seek to quantify species‐specific responses in detection probability and consider how component detection probabilities could influence survey outcomes.     

Traffic noise differentially impacts call types in a Japanese treefrog (Buergeria japonica)

Acoustic noise from automobile traffic impedes communication between signaling animals. To overcome the acoustic interference imposed by anthropogenic noise, species across taxa adjust their signaling behavior to increase signal saliency. As most of the spectral energy of anthropogenic noise is concentrated at low acoustic frequencies, species with lower frequency signals are expected to be more affected. Thus, species with low-frequency signals are under stronger pressure to adjust their signaling behaviors to avoid auditory masking than species with higher frequency signals. Similarly, for a species with multiple types of signals that differ in spectral characteristics, different signal types are expected to be differentially masked. We investigate how the different call types of a Japanese stream breeding treefrog (Buergeria japonica) are affected by automobile traffic noise. Male B. japonica produce two call types that differ in their spectral elements, a Type I call with lower dominant frequency and a Type II call with higher dominant frequency. In response to acoustic playbacks of traffic noise, B. japonica reduced the duration of their Type I calls, but not Type II calls. In addition, B. japonica increased the call effort of their Type I calls and decreased the call effort of their Type II calls. This result contrasts with prior studies in other taxa, which suggest that signalers may switch to higher frequency signal types in response to traffic noise. Furthermore, the increase in Type I call effort was only a short-term response to noise, while reduced Type II call effort persisted after the playbacks had ended. Overall, such differential effects on signal types suggest that some social functions will be disrupted more than others. By considering the effects of anthropogenic noise across multiple signal types, these results provide a more in-depth understanding of the behavioral impacts of anthropogenic noise within a species.     

In Situ Mortality Experiments with Juvenile Sea Bass (Dicentrarchus labrax) in Relation to Impulsive Sound Levels Caused by Pile Driving of Windmill Foundations

Impact assessments of offshore wind farm installations and operations on the marine fauna are performed in many countries. Yet, only limited quantitative data on the physiological impact of impulsive sounds on (juvenile) fishes during pile driving of offshore wind farm foundations are available. Our current knowledge on fish injury and mortality due to pile driving is mainly based on laboratory experiments, in which high-intensity pile driving sounds are generated inside acoustic chambers. To validate these lab results, an in situ field experiment was carried out on board of a pile driving vessel. Juvenile European sea bass (Dicentrarchus labrax) of 68 and 115 days post hatching were exposed to pile-driving sounds as close as 45 m from the actual pile driving activity. Fish were exposed to strikes with a sound exposure level between 181 and 188 dB re 1 µPa².s. The number of strikes ranged from 1739 to 3067, resulting in a cumulative sound exposure level between 215 and 222 dB re 1 µPa².s. Control treatments consisted of fish not exposed to pile driving sounds. No differences in immediate mortality were found between exposed and control fish groups. Also no differences were noted in the delayed mortality up to 14 days after exposure between both groups. Our in situ experiments largely confirm the mortality results of the lab experiments found in other studies.     

Anthropogenic noise affects risk assessment and attention: the distracted prey hypothesis

Many studies have focused on the effects of anthropogenic noise on animal communication, but only a few have looked at its effect on other behavioural systems. We designed a playback experiment to test the effect of noise on predation risk assessment. We found that in response to boat motor playback, Caribbean hermit crabs (Coenobita clypeatus) allowed a simulated predator to approach closer before they hid. Two hypotheses may explain how boat noise affected risk assessment: it masked an approaching predator''s sound; and/or it reallocated some of the crabs'' finite attention, effectively distracting them, and thus preventing them from responding to an approaching threat. We found no support for the first hypothesis: a silent looming object still got closer during boat motor playbacks than during silence. However, we found support for the attentional hypothesis: when we added flashing lights to the boat motor noise to further distract the hermit crabs, we were able to approach the crabs more closely than with the noise alone. Anthropogenic sounds may thus distract prey and make them more vulnerable to predation.     

Experimental Chronic Noise Is Related to Elevated Fecal Corticosteroid Metabolites in Lekking Male Greater Sage-Grouse (Centrocercus urophasianus)

There is increasing evidence that individuals in many species avoid areas exposed to chronic anthropogenic noise, but the impact of noise on those who remain in these habitats is unclear. One potential impact is chronic physiological stress, which can affect disease resistance, survival and reproductive success. Previous studies have found evidence of elevated stress-related hormones (glucocorticoids) in wildlife exposed to human activities, but the impacts of noise alone are difficult to separate from confounding factors. Here we used an experimental playback study to isolate the impacts of noise from industrial activity (natural gas drilling and road noise) on glucocorticoid levels in greater sage-grouse (Centrocercus urophasianus), a species of conservation concern. We non-invasively measured immunoreactive corticosterone metabolites from fecal samples (FCMs) of males on both noise-treated and control leks (display grounds) in two breeding seasons. We found strong support for an impact of noise playback on stress levels, with 16.7% higher mean FCM levels in samples from noise leks compared with samples from paired control leks. Taken together with results from a previous study finding declines in male lek attendance in response to noise playbacks, these results suggest that chronic noise pollution can cause greater sage-grouse to avoid otherwise suitable habitat, and can cause elevated stress levels in the birds who remain in noisy areas.     

Rewiring coral: Anthropogenic nutrients shift diverse coral–symbiont nutrient and carbon interactions toward symbiotic algal dominance

Improving coral reef conservation requires heightened understanding of the mechanisms by which coral cope with changing environmental conditions to maintain optimal health. We used a long‐term (10 month) in situ experiment with two phylogenetically diverse scleractinians (Acropora palmata and Porites porites) to test how coral–symbiotic algal interactions changed under real‐world conditions that were a priori expected to be beneficial (fish‐mediated nutrients) and to be harmful, but non‐lethal, for coral (fish + anthropogenic nutrients). Analyzing nine response variables of nutrient stoichiometry and stable isotopes per coral fragment, we found that nutrients from fish positively affected coral growth, and moderate doses of anthropogenic nutrients had no additional effects. While growing, coral maintained homeostasis in their nutrient pools, showing tolerance to the different nutrient regimes. Nonetheless, structural equation models revealed more nuanced relationships, showing that anthropogenic nutrients reduced the diversity of coral–symbiotic algal interactions and caused nutrient and carbon flow to be dominated by the symbiont. Our findings show that nutrient and carbon pathways are fundamentally “rewired” under anthropogenic nutrient regimes in ways that could increase corals’ susceptibility to further stressors. We hypothesize that our experiment captured coral in a previously unrecognized transition state between mutualism and antagonism. These findings highlight a notable parallel between how anthropogenic nutrients promote symbiont dominance with the holobiont, and how they promote macroalgal dominance at the coral reef scale. Our findings suggest more realistic experimental conditions, including studies across gradients of anthropogenic nutrient enrichment as well as the incorporation of varied nutrient and energy pathways, may facilitate conservation efforts to mitigate coral loss.     

Threshold for onset of injury in Chinook salmon from exposure to impulsive pile driving sounds

The risk of effects to fishes and other aquatic life from impulsive sound produced by activities such as pile driving and seismic exploration is increasing throughout the world, particularly with the increased exploitation of oceans for energy production. At the same time, there are few data that provide insight into the effects of these sounds on fishes. The goal of this study was to provide quantitative data to define the levels of impulsive sound that could result in the onset of barotrauma to fish. A High Intensity Controlled Impedance Fluid filled wave Tube was developed that enabled laboratory simulation of high-energy impulsive sound that were characteristic of aquatic far-field, plane-wave acoustic conditions. The sounds used were based upon the impulsive sounds generated by an impact hammer striking a steel shell pile. Neutrally buoyant juvenile Chinook salmon (Oncorhynchus tshawytscha) were exposed to impulsive sounds and subsequently evaluated for barotrauma injuries. Observed injuries ranged from mild hematomas at the lowest sound exposure levels to organ hemorrhage at the highest sound exposure levels. Frequency of observed injuries were used to compute a biological response weighted index (RWI) to evaluate the physiological impact of injuries at the different exposure levels. As single strike and cumulative sound exposure levels (SEL(ss), SEL(cum) respectively) increased, RWI values increased. Based on the results, tissue damage associated with adverse physiological costs occurred when the RWI was greater than 2. In terms of sound exposure levels a RWI of 2 was achieved for 1920 strikes by 177 dB re 1 μPa(2)?s SEL(ss) yielding a SEL(cum) of 210 dB re 1 μPa(2)?s, and for 960 strikes by 180 dB re 1 μPa(2)?s SEL(ss) yielding a SEL(cum) of 210 dB re 1 μPa(2)?s. These metrics define thresholds for onset of injury in juvenile Chinook salmon.     

Interspecific responses to distress calls in bats (Chiroptera: Vespertilionidae): a function for convergence in call design?

Distress calls were recorded from three sympatric species of pipistrelle bat (Pipistrellus nathusii, P. pipistrellus and P. pygmaeus) in England and Northern Ireland. At foraging sites, we conducted playback experiments, consisting of experimental distress call sequences from each species and control sequences of random noise and sound recorded with no bats present. We measured response by simultaneously recording ultrasound during playbacks and counting the echolocation pulses above a predetermined threshold which were then identified to species. All three species responded to each other's calls. The number of recorded echolocation pulses of all species increased eight-fold, on average, during the playback of distress call sequences compared with the playback of ultrasonic noise, and four-fold compared with the playback of silence. In a separate playback experiment, the number of echolocation pulses of P. pygmaeus increased 14-fold during the playback of distress calls of four endemic species of bat from Madagascar (Emballonura atrata, Myotis goudoti, Miniopterus majori and M. manavi) compared with the playback of silence. This increased response might have been caused by the high calling rates of the Malagasy species. Distress calls of P. nathusii, P. pipistrellus and P. pygmaeus were structurally convergent, consisting of a series of downward-sweeping, frequency-modulated elements of short duration and high intensity with a relatively strong harmonic content. Selection may favour convergence in the structure of distress calls among bat species, if attracting heterospecifics increases the chance of repelling predators by mobbing.     

Nocturnal playbacks reveal hidden differences in singing behaviour between populations of Robin Erithacus rubecula

Capsule Birds subjected to playbacks of conspecific song responded strongly by day in both Wales and Ireland, and strongly at night only in Wales.

Aims To test the responses of Robins to conspecific playbacks between Wales and Ireland and between day and night.

Methods We measured the responses of Robins to conspecific playbacks during daylight, and at night on territories illuminated by streetlights, and compared Robins in Wales and in Ireland.

Results Most Welsh Robins responded to playbacks at night, whereas Irish Robins rarely responded. Both populations responded equally during daylight.

Conclusion We show for the first time that different populations with similar daytime behaviours may differ in nocturnal behaviour. Such population differences in communication behaviour should be taken into account, for example when using acoustic playback to census populations.     

Ready to Fight: Reliable Predictors of Attack in a Cooperatively Breeding,Non‐Passerine Bird

Signal reliability is a major focus of animal communication research. Aggressive signals are ideal for measuring signal reliability because the signal referent – attack or no attack – can be measured unambiguously. Signals of aggressive intent occur at elevated rates in aggressive contexts, predict subsequent aggression by the signaler, and elicit appropriate responses from receivers. We tested the ‘predictive criterion’ in smooth‐billed anis, Crotophaga ani, by broadcasting one of two playback types (‘ahnee’ calls only or ‘ahnee + hoot’ calls), presenting a taxidermic mount, and observing the animals’ behavior. Based on the hypotheses that ‘hoot’ calls and ‘throat‐inflation’ displays signal aggressive intent, we predicted that they would be associated with attack, and that signaling rate would increase over the time period leading up to an attack. Indeed, both hoots and throat‐inflation displays reliably predicted attack. The second prediction, that signaling rate increases in the time leading up to attack, was strongly supported for throat‐inflation displays, which increased over the pre‐attack period in both treatments. Hoots increased over the pre‐attack period in ahnee playbacks but not in ahnee + hoot playbacks. Hierarchical signaling systems are characterized by early, less‐reliable predictors of attack, and later, more reliable predictors of attack. During both natural and simulated interactions, the more‐reliable throat‐inflation display tended to precede the less‐reliable hoot call, suggesting that this signaling system is not hierarchical. In a comparison of 11 putative signals of aggressive intent in birds, the throat‐inflation display had the second highest mutual information (reduction in uncertainty) among visual signals and non‐passerine signals while hoots had below‐average mutual information. Natural observations indicate that both hoots and throat‐inflation displays occur in the context of aggressive between‐group encounters, and hoots also occur during within‐group interactions. Throat‐inflation displays appear to be reliable indicators of aggressive intent, but the function of hoot calls is less clear.