Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

Social cues facilitate habitat selection: American redstarts establish breeding territories in response to song

For migratory songbirds nesting in northern temperate forests, a short breeding season demands that males rapidly establish territories. Because critical insect and vegetation resources are unavailable during spring arrival, we suggest that conspecifics serve as settlement cues for males new to a local population. To test conspecific attraction, we conducted playback experiments with American redstarts Setophaga ruticilla. Experimental results demonstrate that song playbacks strongly attract conspecifics, recruiting an average of 4.2 additional males per plot; adult males new to our sites increased, while yearling males failed to respond. Yearlings arrived 6 to 10 days later than adults, raising the possibility that yearlings responded to songs of early arriving adults rather than to playbacks. Our work indicates that conspecific attraction is an important mechanism for breeding habitat selection in an established population of a migratory forest songbird, but the effect is moderated by age, reproductive experience and arrival timing.     

Conspecific attraction in a grassland bird, the Baird's Sparrow

ABSTRACT.   Territorial songbirds generally use song to defend territories and attract mates, but conspecific song may also serve as a cue to attract other male songbirds to a breeding site. Although known to occur in some colonial and forest-associated species, only recently have investigators examined conspecific attraction in grassland species. We used a playback experiment to examine the possible role of conspecific attraction for males searching for potentially suitable breeding habitat in a grassland specialist, the Baird's Sparrow ( Ammodramus bairdii ). Experimental playback plots and control plots with similar landscape and vegetation characteristics were established at two sites in North Dakota. Baird's Sparrows colonized three of six experimental plots and none of six control plots. Males on experimental plots established territories adjacent to the playback stations and were sometimes observed counter-singing with the playback of conspecific songs. Vegetation characteristics were similar on all study plots, and did not explain differences in bird density on our treatment plots. Although we found that playback of conspecific songs attracted male Baird's Sparrows to previously unoccupied, potentially suitable habitat, further experiments are needed to examine the importance of conspecific attraction relative to other cues that birds may use, such as vegetation features. The conservation and management implications of conspecific attraction are not completely understood, but the presence of conspecifics should be considered as a potential cue in habitat selection by all species of birds.     

Social information trumps vegetation structure in breeding-site selection by a migrant songbird

To maximize fitness, organisms must assess and select suitable habitat. Early research studying birds suggested that organisms consider primarily vegetation structural cues in their habitat choices. We show that experimental exposure to singing in the post-breeding period provides a social cue that is used for habitat selection the following year by a migrant songbird, the black-throated blue warbler (Dendroica caerulescens). Our experimental social cues coerced individuals to adopt territories in areas of very poor habitat quality where individuals typically do not occur. This indicates that social information can override typical associations with vegetation structure. We demonstrate that a strong settlement response was elicited because post-breeding song at a site is highly correlated with reproductive success. These results constitute a previously undocumented, but highly parsimonious mechanism for the inadvertent transfer of reproductive (public) information from successful breeders to dispersers. We hypothesize that post-breeding song is a pervasive and reliable cue for species that communicate vocally, inhabit temporally autocorrelated environments, produce young asynchronously and/or abandon territories after reproductive failure.     

Experimental Tests of the Function and Flexibility of Song Consistency in a Wild Bird

Measures of bird song that capture aspects of motor performance, such as consistency, have become a major focus in understanding sexual selection on song. Despite accumulating evidence that consistency is related to reproductive success in many species, the relative importance of male–male interactions and female–male interactions is still unclear. We studied the function and flexibility of song consistency and song rate in common yellowthroat warblers (Geothlypis trichas). A previous study of this population found that song consistency—measured as the amount of variability within a bout of songs—was positively correlated with the likelihood of siring extrapair young. In this study, we conducted two experiments aimed at testing (1) the role of song consistency and rate in mediating male–male and male–female interactions and (2) whether song effort is flexibly adjusted to changes in social context. In the first experiment, we simulated a male territorial intrusion with song playbacks that varied in consistency and rate; focal males responded aggressively to playbacks, but their response did not differ with playback consistency or rate. In the second experiment, we presented focal males with a taxidermic female mount and female vocalizations; focal males approached the speaker, but continued to sing and did not perform the aggressive rattle vocalization observed during male encounters. Immediately after the simulated female encounter, focal males increased in song consistency. Taken together, our results are most consistent with the hypothesis that song consistency in common yellowthroats is primarily a female‐directed signal that is actively adjusted in response to rapidly changing social conditions.     

Avian Mobbing Response is Restricted by Territory Boundaries: Experimental Evidence from Two Species of Forest Warblers

Predator mobbing has been viewed as an adaptation to reduce the risk of predation, however, factors influencing mobbing behaviour are still debated. We report on the results of an experiment with Dendroica caerulescens and Dendroica virens designed to determine (1) whether mobbing response by forest songbirds during the breeding season is restricted by territory boundaries, (2) the distance songbirds will move in response to anti‐predator mobbing calls, and (3) whether reproductive status, age, and time of the breeding season determine the distance moved to mob. We did not detect an effect of reproductive status, age, or time of breeding season on the distance moved by birds to mob. All birds responded to the mobbing playback within their territory (defined by territorial defence in relation to specific song playbacks). The maximum distance moved within a territory to engage in mobbing ranged from 25 to 175 m ( = 72 ± 6 m). Three of 37 birds responded to playbacks outside their territory boundaries. In all three cases, maximum movement distances outside territories were short (25 m). Thus, for two species of warblers, mobbing is highly constrained by territory boundaries during the breeding season. This finding is congruent with arguments that mobbing is primarily a selfish behaviour, at least with respect to conspecifics. Our results also provide support for the ‘move‐on’ hypothesis.     

The Return of the Intruder: Immediate and Later Effects of Different Approach Distances in a Territorial Songbird

Male songbirds often maintain territories throughout the breeding season, and one of the main functions of song is to deter invaders. Therefore, the distance of an unknown singing rival should play a crucial role within territorial singing interactions of males. This distance is expected to be assessed as more threatening the closer the rival approaches. Here, we tested this assumption by conducting nocturnal playbacks from two different distances in territorial Common nightingales (Luscinia megarhynchos). Immediate vocal responses of birds were examined by analysing changes in song structure as well as temporal response features. The next morning, follow‐up playbacks from an intermediate distance allowed us to investigate longer‐lasting effects of nocturnal playbacks. We found that the distance of a simulated rival had an effect on both immediate and later vocal responses of territorial male nightingales with different song parameters being affected during nocturnal and diurnal singing. This indicates that birds perceive intruder distances and adjust their response behaviour both immediately and in later interactions.     

Malarial parasites as geographical markers in migratory birds?

We tested the hypothesis that malarial parasites (Plasmodium and Haemoproteus) of black-throated blue warblers (Dendroica caerulescens) provide sufficient geographical signal to track population movements between the warbler's breeding and wintering habitats in North America. Our results from 1083 warblers sampled across the species' breeding range indicate that parasite lineages are geographically widespread and do not provide site-specific information. The wide distribution of malarial parasites probably reflects postnatal dispersal of their hosts as well as mixing of breeding populations on the wintering range. When compared to geographically structured parasites of sedentary Caribbean songbirds, patterns of malarial infections in black-throated blue warblers suggest that host-malaria dynamics of migratory and sedentary bird populations may be subject to contrasting selection pressures.     

Multiple Plumage Ornaments as Signals of Intrasexual Communication in Golden‐Winged Warblers

Avian plumage represents some of the greatest diversity in integument coloration of all animals. Plumage signals are diverse in function, including those that allow for assessing potential mates or the mitigation of agonistic interactions between rivals. Many bird species possess multiple ornamental traits that have the potential to serve as multiple or redundant signals. For example, male golden‐winged warblers (Vermivora chrysoptera) have brilliant carotenoid‐based yellow crowns, melanin‐based black throats, and structurally based white patches on their outer tail feathers. Using a correlative approach, we investigated whether plumage ornaments have the potential to reliably signal ability to acquire higher quality territory, aggressive response to simulated territorial intrusions, and reproductive success. We found that both crown chroma and tail brightness were significantly related to habitat quality and aggression; more ornamented birds held territories with higher quality habitat and were less aggressive toward simulated conspecific stimuli. Older birds sang less threatening songs than younger birds and were more likely to sing their mate attraction song type (type 1) rather than songs typically reserved for agonistic interactions (type 2). Finally, despite our previous research demonstrating that habitat strongly predicts reproductive success in this warbler population, we found no evidence of a direct link between ornamentation and reproductive success. Overall, these data suggest that younger males, and those with lower quality ornaments, compensate with more aggressive behaviors. Additional research is needed to investigate the dynamics between behavioral traits and ornaments to better understand complex signaling and how golden wing signals function in conspecific interactions (male–male interactions and mate‐choice).     

Hostile encounters among spruce-woods warblers (Dendroica: Parulidae)

Intraspecific interactions were initiated with high frequency by four species of sympatric warblers (Dendroica: Parulidae) early in the breeding season. They declined during the incubation and nestling stages and then increased during the fledging stage in two species, the black-throated green (Dendroica virens) and yellow-rumped (D. coronata) warblers. Interspecific interactions occurred infrequently early in the season, but increased during the incubation and nestling stages in the black-throated green and blackburnian (D. fusca) warblers and during the fledging stage in the magnolia warbler (D. magnolia). Fights and chases predominated over supplanting attacks in both intraspecific and interspecific encounters. Male black-throated green and yellow-rumped warblers initiated hostile encounters more frequently than did their females, but male and female blackburnian and magnolia warblers did not differ in this regard. A somewhat equivocal interspecific social dominance hierarchy occurred: magnolia > black-throated green > yellow-rumped > blackburnian. Reversals to this trend were frequent and may have resulted from the strong defences initiated by these species in certain parts of their territories, particularly about their nests.     

Song learning by prairie warblers: When,where, and from whom

Songbirds of many species acquire their songs by imitating the songs of conspecific singers. Conclusive evidence of such imitation comes from controlled laboratory studies, but such studies do not reveal when and where songbirds learn their songs under natural conditions. To determine the timing and location of song learning in a population of prairie warblers, we compared the songs of yearling prairie warblers of known hatching location to the songs of other birds in the yearlings' natal and first breeding areas. The comparisons yielded a likely model song (and model singer) for each of the song types used by the focal yearlings. We supplemented our findings from the song comparisons with inferences drawn from an analysis of local geographic variation in songs. This analysis revealed that shared song types showed no tendency to be geographically clustered within the study area. Taken together, our data suggest that prairie warblers learn their songs during the hatch year, at locations somewhat distant (mean distance 1,437 m) from their natal site, most likely as birds wander about during the post-fledging period.     

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.     

MICRODIALECT AND GROUP SIGNATURE IN THE SONG OF THE SKYLARK ALAUDA ARVENSIS

ABSTRACT

The Skylark Alauda arvensis is a territorial species of open landscape in which pairs settle in stable and adjacent territories during the breeding season. Due to the heterogeneity of the habitat, territories are gathered in patches spaced by a few kilometres, in which each male produces very long and complex flight songs as a part of the territorial behaviour. We showed that, in a given patch, all the males (neighbours) share some particular sequences of syllables in their songs, whereas males settled in different patches (strangers) have almost no sequences in common. Such a phenomenon is known as microdialect. To test the hypothesis that these shared sequences support a group signature, we made playback experiments with “chimeric” signals: songs of strangers where the sequences shared by neighbours were artificially inserted. Behavioural responses to playbacks indicated a neighbour-stranger discrimination consistent with the dear enemy phenomenon, i.e. a reduced aggression toward neighbours compared to strangers. Furthermore, the same level of responses, observed when a “chimeric” song and a neighbour song were broadcast, indicated that shared sequences are recognised and identified as markers of the neighbourhood identity.     

Sexual differentiation and seasonal variation in response to conspecific and heterospecific acoustic signals

Interspecific territoriality is frequently reported between closely related species; however, few studies have demonstrated interspecific territoriality between distantly related species living in sympatry. We conducted playback experiments to investigate territorial behaviour in male and female White‐bellied Wrens (Uropsila leucogastra) in response to simulated conspecific and heterospecific intruders during the breeding and non‐breeding seasons. We explored whether heterospecific songs of the Happy Wren (Pheugopedius felix), a distantly related species and ecological competitor, elicited antagonistic responses from focal White‐bellied Wrens, and whether such responses differed between the sexes. We also examined whether male and female responses to conspecific and heterospecific rivals varied with season. We found that male White‐bellied Wrens always responded to conspecific song, and responded significantly more to heterospecific song compared to a control stimulus (Tropical Parula, Setophaga pitiayumi). In contrast, although female White‐bellied Wrens responded strongly to conspecific song, their response to heterospecific song did not differ significantly from the control stimulus. The proportion of males that responded to heterospecific songs and the proportion of females that responded to conspecific songs varied seasonally, showing significantly lower responses during the breeding season. The intense responses of male White‐bellied Wrens to playback of heterospecific songs suggest that they recognise ecological competitors based on their vocal signals. Furthermore, the decrease in agonistic interactions during the breeding season is in line with the hypothesis that aggressive behaviour may be detrimental to reproductive and parental activity, and the hypothesis that heterospecific animals pose less of a threat during the breeding season.     

Duetting behavior in a Neotropical ovenbird: sexual and seasonal variation and adaptive signaling functions

Duetting is a collective behavior and might have multiple functions, including joint territory defense and mate guarding. An important step toward understanding the adaptive function of bird song is to determine if and how singing behavior varies seasonally. However, seasonal patterns for duetting species are different from the pattern described for species in which only the male sings, because song function may vary according to sex, singing role (initiator vs responder) and level of duet organization (individual vs pair). We investigated whether patterns of seasonal variation in duetting depends on these factors, which would suggest different interpretations of song function. We studied social pairs of a Neotropical bird species (rufous hornero Furnarius rufus) for seven consecutive months, recording vocal and territorial behaviors. Overall, partners coordinated 61% of their songs into duets and many song traits (song initiation rate, song output and duet rate) peaked in territorial contexts. Males engaged in territorial interactions with strangers more often, initiated more songs, and answered proportionately more of their partners’ songs than females. Male song initiation rate peaked during the pre‐ and post‐breeding stages, whereas females initiated more songs during the non‐breeding season. Both sexes answered partner songs faster and at higher rates during the pre‐breeding and female fertile stages. Partners duetted at a higher rate during the pre‐ and post‐breeding stages. Finally, song initiation rates and duet rate, but not song answering rates, correlated with frequency of territorial interactions with strangers. Although our findings indicate that song function may vary with sex, singing role and level of duet organization, our results suggest that in general duet functions to defend common territories and as a mutual mate guarding strategy in the rufous hornero.     

Territorial aggression can be sensitive to the status of heterospecific intruders

Territorial animals are known to be able to differentiate between intruding individuals posing a low or high threat and adjust their aggressive response accordingly. However, plastic territorial aggression based on recognising individuals with different attributes is typically assumed to be relevant only in the context of conspecific interactions. In this study, we investigated territorial aggression of neotropical cichlid fish in their natural habitat to assess whether responses to different types of individuals of another species can also be plastic. We show that arrow cichlids (Amphilophus zaliosus) adjusted their territorial aggression regarding the status of heterospecific intruders: breeding individuals of Amphilophus astorquii received a lower level of aggression than non-breeders. The same pattern was also found for the two different types of A. astorquii individuals intruding into conspecific territories. These results suggest that heterospecific individuals should not be ignored when considering selection pressures shaping plasticity of aggressive behaviour in territorial animals.     

On the song resumption, polyterritorial behaviour and their population context in the Sedge Warbler Acrocephalus schoenobaenus

Mating tactics in species with facultative polygyny seem to be very flexible and to depend on local environmental conditions. We analysed the habitat and population contexts of territorial behaviour, associated with polygyny, in a population of Sedge Warblers Acrocephalus schoenobaenus inhabiting natural wetlands. Nearly one-third of all breeding males (46 individuals) resumed song after completing their mating with the first female, in order to attract another one. Resuming males were usually the earliest arrivals. There was a continuity in the territorial behaviour between resuming song on first territory and polyterritorial behaviour. Fifty-nine percent of resuming males set up second territories, clearly separated from the first. The second territory was usually located close to the primary female activity area. The quality of the first and second territories were correlated, with second territories being significantly inferior. However, their quality was not significantly different from the territories of non-breeding males. The quality of the second territories was also negatively related to the distance from the first territories. Polyterritorialism was influenced by population numbers: the higher the number of territorial males, the lower the number of second territories settled and the lower their average quality. Although the polygyny frequency in the studied population was very low, polyterritorial males were significantly more likely to be polygynous. We concluded that polygyny frequency can be significantly influenced by population numbers, which might be one of the main factors responsible for the variability in the mating system in this species.     

Escalation of aggressive vocal signals: a sequential playback study

Rival conspecifics often produce stereotyped sequences of signals as agonistic interactions escalate. Successive signals in sequence are thought to convey increasingly pronounced levels of aggressive motivation. Here, we propose and test a model of aggressive escalation in black-throated blue warblers, presenting subjects with two sequential and increasingly elevated levels of threat. From a speaker outside the territorial boundary, we initiated an interaction (low-threat level), and from a second speaker inside the territory, accompanied by a taxidermic mount, we subsequently simulated a territorial intrusion (escalated threat level). Our two main predictions were that signalling behaviours in response to low-threat boundary playback would predict signalling responses to the escalated within-territory threat, and that these latter signalling behaviours would in turn reliably predict attack. We find clear support for both predictions: (i) specific song types (type II songs) produced early in the simulated interaction, in response to boundary playback, predicted later use of low-amplitude ‘soft’ song, in response to within-territory playback; and (ii) soft song, in turn, predicted attack of the mount. Unexpectedly, use of the early-stage signal (type II song) itself did not predict attack, despite its apparent role in aggressive escalation. This raises the intriguing question of whether type II song can actually be considered a reliable aggressive signal. Overall, our results provide new empirical insights into how songbirds may use progressive vocal signalling to convey increasing levels of threat.     

Does interspecific competition affect territorial distribution of birds? A long‐term study on Siberian Phylloscopus warblers

Seven species of Phylloscopus warblers breed in the middle taiga subzone of Central Siberia. We studied their territorial distribution over 18 breeding seasons between 1978 and 1996. Phylloscopus inornatus , the most abundant breeding bird species in the study area, breeds in dense clusters of territories. In some habitats, these clusters are very unstable from year to year, providing an excellent opportunity to study the influence of P. inornatus on the territorial distribution of other species. We present a method for quantifying decreases in a species' breeding density caused by the presence of a competing congener. Analysing all possible combinations of species, we found reciprocal territorial avoidance between P. inornatus and P. proregulus as well as between P. inornatus and P. collybita . This is consistent with the hypothesis that territorial avoidance occurs between the species of greatest morphological similarity. The occurrence of territorial avoidance is not explained by habitat characteristics such as vegetation cover and productivity. However low productivity may be necessary for territorial avoidance to occur. An alternative explanation for the observed territorial separation of some species, derived from the Fretwell-Lucas model of habitat occupancy, is examined and rejected. Causes and implications of clustering behaviour of P. inornatus are also discussed.     

Stable isotope ratios indicate that body condition in migrating passerines is influenced by winter habitat

Although predicted some time ago, there has been little success in demonstrating that the overall fitness of migratory birds depends on the combined influences of their experiences over all seasons. We used stable carbon isotope signatures (delta13C) in the claws of migrating black-throated blue warblers Dendroica caerulescens to infer their wintering habitats and investigated whether winter habitat selection can be linked to condition during migration. Resident bird species with low delta13C corresponded to selection of more mesic habitats, and migrating birds with low delta13C were in better condition than conspecifics with higher delta13C signatures. These findings concur with empirical observations on the wintering grounds, where dominants (mostly males) tend to exclude subordinates from mesic areas (considered to be high-quality habitats). We believe that variation in condition during migration may be one of the key factors determining differences in arrival times and condition at the breeding areas, which in turn have a major influence on reproductive success.