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  1. Referential alarm calls that denote specific types of dangers are common across diverse vertebrate lineages. Different alarm calls can indicate a variety of threats, which often require specific actions to evade. Thus, to benefit from the call, listeners of referential alarm calls must be able to decode the signaled threat and respond to it in an appropriate manner. Yellow warblers ( Setophaga petechia ) produce referential “seet” calls that signal to conspecifics the presence of nearby obligate brood parasitic brown-headed cowbirds ( Molothrus ater ), which lay their eggs in the nests of other species, including yellow warblers. Our previous playback experiments have found that red-winged blackbirds ( Agelaius phoeniceus ), a species also parasitized by brown-headed cowbirds, eavesdrop upon and respond strongly to yellow warbler seet calls during the incubation stage of breeding with aggression similar to responses to both cowbird chatters and predator calls. To assess whether red-winged blackbird responses to seet calls vary with their own risk of brood parasitism, we presented the same playbacks during the nestling stage of breeding (when the risk of brood parasitism is lower than during incubation). As predicted, we found that blackbirds mediated their aggression toward both cowbird chatter calls and the warblers’ anti-parasitic referential alarm calls in parallel with the low current risk of brood parasitism during the nestling stage. These results further support that red-winged blackbirds flexibly respond to yellow warbler antiparasitic referential calls as a frontline defense against brood parasitism at their own nests. 
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  2. Many avian species are negatively impacted by obligate avian brood parasites, which lay their eggs in the nests of host species. The yellow warbler (Setophaga petechia), which is host to the brood-parasitic brown-headed cowbird (Molothrus ater), represents one of the best-replicated study systems assessing antiparasitic host defenses. Over 15 prior studies on yellow warblers have used model-presentation experiments, whereby breeding hosts are exposed to models of brown-headed cowbirds or other nest threats, to test for anti-parasitic defenses unique to this species. Here we present results from our own quasi-replication study of the yellow warbler/brown-headed cowbird system, which used a novel design compared to previous experiments by pivoting to conduct acoustic playback treatments only, rather than presenting visual models with or without calls. We exposed active yellow warbler nests to playbacks of brown-headed cowbird chatters (brood parasite), blue jay (Cyanocitta cristata; nest predator) calls, conspecific “seet” calls (a referential alarm call for brood parasitism risk), conspecific “chip” calls (a generic alarm call), or control wood thrush (Hylocichla mustelina; harmless heterospecific) songs during the incubation stage. Similar to previous studies, we found that female yellow warblers seet called more frequently in response to playbacks of both brood parasitic chatter calls and conspecific seet calls whereas they produced more chip calls in response to the playback of nest predator calls. In contrast, female yellow warblers approached all playbacks to similar distances, which was different from the proximity patterns seen in previous studies. Our study demonstrates the importance of both replicating, and also pivoting, experimental studies on nest defense behaviors, as differences in experimental design can elicit novel behavioral response patterns in the same species. 
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  3. Abstract

    Vocal signals mediate social relationships, and among networks of territorial animals, information is often shared via broadcast vocalizations. Anthropogenic noise may disrupt communication among individuals within networks, as animals change the way they vocalize in noise. Furthermore, constraints on signal transmission, including frequency masking and distance, may affect information exchange following a disruption in social networks. We tested the hypothesis that signaling interactions within networks of breeding male and female house wrens (Troglodytes aedon) depend on distance, ambient noise, and receiver nesting stage. We used playback experiments to simulate territorial intrusions with and without noise playbacks on the territories of established males and simultaneously recorded the vocal responses of neighbors. To examine whether intrusions impacted interactions between males, we used randomization tests to determine whether treatment, distance, noise, or nesting stage affected vocal coordination between challenged and neighboring males. We also quantified singing patterns to explore whether intrusions on territories of challenged males affected singing by males and females on neighboring territories. Males sang at the lowest rates and were less likely to overlap songs with the challenged male when their partner was laying, compared to males during early and late nesting stages. Noise and distance did not affect vocal coordination or male singing rates. Fewer females sang during the intruder‐only treatment compared to the control and intrusions with noise. Added noise in the territories of challenged males may have masked signals, and as a result, females only changed their behavior during the intruder‐only treatment. Our results suggest that the fertility of breeding partners may be more important to males than short‐term changes on rival male territories. Elevated noise did little to alter male responses to threats within networks. Females appeared to eavesdrop on interactions involving neighboring males, but noise may have prevented detection of their interactions.

     
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  4. Abstract

    Defending offspring incurs temporal and energetic costs and can be dangerous for the parents. Accordingly, the intensity of this costly behavior should reflect the perceived risk to the reproductive output. When facing costly brood parasitism by brown‐headed cowbirds (Molothrus ater), where cowbirds lay eggs in heterospecific nests and cause the hosts to care for their young, yellow warblers (Setophaga petechia) use referential “seet” calls to warn their mates of the parasitic danger. Yellow warblers of both sexes produce this call only in response to cowbirds or seet‐calling conspecifics. Seet calls are mainly produced during the laying and incubation stages of breeding, when risk of brood parasitism is highest, rather than during the nestling stage. On the other hand, general alarm calls (chips) are produced throughout the nesting cycle and are also used in conspecific interactions unrelated to nesting. We hypothesized that context shapes responses prior to breeding as well, such that yellow warblers without a mate and active nest would be less likely to respond to playbacks that simulate brood parasitism risk. To test this hypothesis, we presented playbacks of two nest threats, cowbirds (brood parasite) and blue jays (Cyanocitta cristata; nest predator), on territories of unmated male warblers (unpaired) and male warblers with a known mate (paired). We found that unpaired males were unresponsive toward playbacks indicating nest threats, whereas paired males were significantly more aggressive and vocal toward these playbacks compared to control playbacks. However, both paired and unpaired males were vocally responsive toward chip calls, which are informative for males regardless of pairing status. Male yellow warblers appear to adjust their responses during the earliest stages of breeding depending on the contextual relevance of specific threat stimuli, and together with prior studies, our work further supports that referential seet calls are associated with stage‐specific risk of brood parasitism.

     
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  5. Yellow warblers ( Setophaga petechia ) use referential ‘seet’ calls to warn mates of brood parasitic brown-headed cowbirds ( Molothrus ater ). In response to seet calls during the day, female warblers swiftly move to sit tightly on their nests, which may prevent parasitism by physically blocking female cowbirds from inspecting and laying in the nest. However, cowbirds lay their eggs just prior to sunrise, not during daytime. We experimentally tested whether female warblers, warned by seet calls on one day, extend their anti-parasitic responses into the future by engaging in vigilance at sunrise on the next day, when parasitism may occur. As predicted, daytime seet call playbacks caused female warblers to leave their nests less often on the following morning, relative to playbacks of both their generic anti-predator calls and silent controls. Thus, referential calls do not only convey the identity or the type of threat at present but also elicit vigilance in the future to provide protection from threats during periods of heightened vulnerability. 
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