Testing the conspecific attraction hypothesis with Dupont's Larks,a resident species of songbird in central Spain

Some birds use social cues, such as the presence of conspecifics, when selecting breeding habitat. This phenomenon, known as conspecific attraction, has been well‐documented in migratory species, but has not been assessed for resident species of birds. We used Dupont's Larks (Chersophilus duponti) as a model species to determine if conspecific attraction plays a role in habitat selection by resident species of birds. At our study site in Soria province in central Spain, we monitored two potential habitat patches and one managed site where management actions had provided apparently suitable habitat. At each site, we broadcast recordings of the songs and calls of male Dupont's Larks, and monitored their presence during the breeding season and dispersal period in 2018 using automated recorders and field surveys. No birds were attracted to our study sites. Our results suggest that management of patches of suitable habitat should occur close to areas (within 1 km) already occupied by Dupont's Larks to encourage natural colonization because, based on our results, playback of conspecific vocalizations may not attract the species to new breeding areas. However, additional studies are needed before drawing conclusions about the effectiveness of conspecific attraction for this and other resident species of birds.     

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.     

Alarm calls modulate the spatial structure of a breeding owl community

Animals should continuously assess the threat of predation. Alarm calls inform on predation risk and are often used as cues to shape behavioural responses in birds and mammals. Hitherto, however, the ecological consequences of alarm calls in terms of organization of animal communities have been neglected. Here, we show experimentally that calls of a resident nocturnal raptor, the little owl Athene noctua, triggered a response in terms of breeding habitat selection and investment in current reproduction in conspecifics and heterospecifics. Little owls preferred to settle in territories where calls of conspecifics, irrespective of their type (i.e. alarm versus contact calls), were broadcasted, indicating that either conspecific attraction exists or calls are interpreted as foreign calls, eliciting settlement as a mode of defence against competitors. Also, we found that little owls seemed to invest more in current reproduction in safe territories as revealed by conspecific calls. Innovatively, we reported that a second owl species, the migratory scops owl Otus scops, preferred to breed in safe territories as indicated by little owls' calls. These results evidence that the emission of alarm calls may have, apart from well-known behavioural effects, ecological consequences in natural communities by inducing species-specific biases in breeding habitat selection. This study demonstrates a previously unsuspected informative role of avian alarm calls which may modulate the spatial structure of species within communities.     

Effects of conspecific and heterospecific residents on patterns of immigration in two species of voles

Studies with birds have shown that presence and density of resident conspecifics and heterospecifics can influence patterns of habitat selection. There have been few studies on the effects of social cues on rates of immigration in mammals. We report results from a long-term live trapping study of immigration in two species of voles, Microtus ochrogaster Wagner, 1842 and Microtus pennsylvanicus Ord, 1815, in bluegrass habitat in east-central Illinois, USA. We compare immigration into control sites from which no individuals of either species were removed with immigration into experimental sites from which either conspecifics or heterospecifics were removed. We focus on characteristics of immigrants and rates of immigration in relation to density in destination habitats. Within each species, immigrants into control and removal sites were similar with respect to body mass and reproductive condition, indicating no major differences in the physical condition of immigrants into sites with established populations and sites without established populations. For both species, density of conspecifics at a site positively influenced rate of immigration at that site. Density of heterospecifics at destination sites did not significantly influence rate of immigration for either species. These results suggest that site selection by dispersing M. ochrogaster and M. pennsylvanicus is characterized by conspecific attraction.     

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.     

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.     

Mechanisms of colony selection by first‐year Lesser Kestrels Falco naumanni

In colonial species, first‐time breeders may use the number of settled conspecifics in colony selection, but such a relationship is confused by the correlation between colony size and nest‐site availability. To distinguish conspecific attraction from neutral colony selection, we experimentally increased nest availability for first‐year Lesser Kestrels Falco naumanni, allowing us to dissociate the number of vacant nest‐sites from colony size at the arrival time of first‐year birds. Under natural conditions, the number of first‐year birds settling was positively correlated with both the number of philopatric and the total number of breeding pairs (colony size) already settled. However, the probability of occupation of experimentally manipulated nests by first‐year birds was independent of colony size. In experimental colonies, the number of first‐year birds settling was positively correlated with the number of manipulated nest‐sites but not with the number of conspecifics. Overall, these results support a neutral colony selection by first‐year Lesser Kestrels based on nest‐site 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.     

Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection. Received: 14 July 1997 / Accepted: 17 November 1997     

Behavioural constraints and conservation biology: Conspecific attraction and recruitment

Wildlife managers have devoted considerable time and research to determine how animals judge the suitability and quality of their habitat. These analyses typically center on habitat characteristics and often produce equivocal results. An additional method of habitat assessment is to examine the extent to which animals, and particularly birds, use conspecifics as cues to establishing breeding and feeding territories. Conspecifics can be indicators of habitat quality, or they might intrinsically affect reproductive success. Here we discuss the implications of conspecific attraction for biologists who wish to conserve endangered species that are living in fragmented habitats.     

Assessing aural and visual cueing as tools for seabird management

Social attraction, that is, mimicking of active and productive colonies via audio playback of calls of breeding conspecifics and the use of decoys, is commonly used to attract birds to newly established or restored breeding sites. However, little is known about the relative importance of aural versus visual cues for identify nesting areas. Such information is important for design and evaluation of management protocols. We studied the effectiveness of decoys (visual cues) and playbacks (audio cues) as methods for restoring a colony of common terns (Sterna hirundo) at Muskeget Island, Massachusetts, USA. We used a 2-year, crossover experiment with 3 treatment areas: audio and visual, audio only, and visual only. We reversed treatment areas in the second year to control for previous nesting area or substrate preference. In both years, nests were built 9–101 m downwind of loudspeakers. There was no overlap in areas used for nesting between years and no nests were built within decoy plots in either year. Behavioral observations showed that birds responded to decoys only when within range of sound treatments. Conspecific vocalizations appear to be important proximate cues for seabird colony site selection and should be given priority in management protocols using social attraction. © 2011 The Wildlife Society.     

Prospecting enhances breeding success of first-time breeders in the great cormorant, Phalacrocorax carbo sinensis

In many species of colonial seabirds, young birds visit colonies in the years before they start breeding. This prospecting behaviour may allow them to obtain information that could enhance their future breeding success. We examined the reproductive consequences of prospecting behaviour in the colonial great cormorant, and found support for this idea. New breeders that had been prospecting actively in the previous year obtained breeding sites of higher quality (i.e. closer to sites where conspecifics had fledged young in the previous year) and had higher breeding success than those that had been less active. Prospecting occurred mostly late in the breeding season, and coincided with the time when the majority of the eggs had hatched but before the chicks started fledging, that is, when breeding success in the colony reflected habitat suitability. These results are thus consistent with the use of conspecific reproductive performance as a cue for the quality of a breeding habitat as expected from the 'performance-based conspecific attraction hypothesis'. Copyright 1999 The Association for the Study of Animal Behaviour.     

Spatial patterns of the frog Oophaga pumilio in a plantation system are consistent with conspecific attraction

The conspecific attraction hypothesis predicts that individuals are attracted to conspecifics because conspecifics may be cues to quality habitat and/or colonists may benefit from living in aggregations. Poison frogs (Dendrobatidae) are aposematic, territorial, and visually oriented—three characteristics which make dendrobatids an appropriate model to test for conspecific attraction. In this study, we tested this hypothesis using an extensive mark‐recapture dataset of the strawberry poison frog (Oophaga pumilio) from La Selva Biological Station, Costa Rica. Data were collected from replicate populations in a relatively homogenous Theobroma cacao plantation, which provided a unique opportunity to test how conspecifics influence the spatial ecology of migrants in a controlled habitat with homogenous structure. We predicted that (1) individuals entering a population would aggregate with resident adults, (2) migrants would share sites with residents at a greater frequency than expected by chance, and (3) migrant home ranges would have shorter nearest‐neighbor distances (NND) to residents than expected by chance. The results were consistent with these three predictions: Relative to random simulations, we observed significant aggregation, home‐range overlap, and NND distribution functions in four, five, and six, respectively, of the six migrant–resident groups analyzed. Conspecific attraction may benefit migrant O. pumilio by providing cues to suitable home sites and/or increasing the potential for social interactions with conspecifics; if true, these benefits should outweigh the negative effects of other factors associated with aggregation. The observed aggregation between migrant and resident O. pumilio is consistent with conspecific attraction in dendrobatid frogs, and our study provides rare support from a field setting that conspecific attraction may be a relevant mechanism for models of anuran spatial ecology.     

Wood warblers copy settlement decisions of poor quality conspecifics: support for the tradeoff between the benefit of social information use and competition avoidance

Social information use in songbird habitat selection commonly involves a conspecific attraction strategy. Individuals copy the breeding‐site choices of conspecifics, that is, bias their own settlement decisions towards sites (tracts of spatially limited habitat with similar structure) already occupied by others. In order to be adaptive, social information use has to be discriminative. Especially the decisions of good quality individuals, i.e. measuring high at observable fitness correlates, should be copied more frequently than those of poor quality individuals. It is unknown, however, whether songbirds discriminatively use conspecific presence by evaluating the quality of information providers in habitat selection. We experimentally tested whether wood warblers Phylloscopus sibilatrix selectively copied settlement decisions of conspecifics in relation to the quality of observed individuals. We also tested whether the use of social cues was influenced by the population density at a particular site in the preceding year. We found that wood warblers selectively used intraspecific social information, but in a pattern opposite to that expected based on existing hypotheses. Wood warblers copied breeding‐site choices of poor quality conspecifics and despite temporary attraction to sites where the presence of good quality individuals was simulated, they did not ultimately settle near these individuals. Population density in the preceding year did not influence settlement patterns. We argue that when making settlement decisions, wood warblers assessed the expected level of local intraspecific competition and selectively copied breeding‐site choices of conspecifics or refused to settle, depending on competitive abilities of observed individuals. This adds a novel aspect to the patterns and processes of social information use proposed thus far, and provides support for the predicted negative effect of intraspecific competition on benefit of information. Moreover, it seems that habitat selection in wood warblers is a complex decision‐making process, in which initial decisions are adjusted after acquiring more accurate information. Synthesis Social information use in songbird habitat selection commonly involves copying the breeding‐site choices of conspecifics (so‐called conspecific attraction). To be adaptive, this strategy has to be discriminative, but almost no empirical studies have tested this assertion. Our study shows that birds may selectively use social information by copying settlement decisions of poor quality conspecifics, but avoid settling near good quality individuals, likely because of their high competitive abilities. This decision‐making pattern supports the predicted, yet not experimentally tested, tradeoff between information value and cost of competition in social information use. Our study highlights also that the use of social cues in settlement decisions may be both positively and negatively biased.     

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.     

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.     

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.     

Conspecific attraction and nest site selection in a nomadic species,the zebra finch

Conspecific nesting density affects many aspects of breeding biology, as well as habitat selection decisions. However, the large variations in breeding density observed in many species are yet to be fully explained. Here, we investigated the settlement patterns in a colonial species with variable breeding density and where resource distribution could be manipulated. The zebra finch, Taeniopygia guttata, is a classic avian model in evolutionary biology but we know surprisingly very little about nest site selection strategies and nesting densities in this species, and in fact, in nomadic species in general. Yet, important determinants of habitat selection strategies, including temporal predictability and breeding synchrony, are likely to be different in nomadic species than in the non‐nomadic species studied to date. Here, we manipulated the distribution of nesting sites (by providing nest boxes) and food patches (feeders) to test four non‐exclusive habitat selection hypotheses that could lead to nest aggregation: 1) attraction to resources, 2) attraction to breeding conspecifics, and 3) attraction to successful conspecifics and 4) use of private information (i.e. own reproductive success on a site). We found that wild zebra finches used conspecific presence and possibly reproductive success, to make decisions over where to locate their nests, but did not aggregate around water or food within the study areas. Moreover, there was a high degree of inter‐individual variation in nesting density preference. We discuss the significance of our results for habitat selection strategy in nomadic species and with respect to the differential selection pressures that individuals breeding at different densities may experience.     

Inter-individual variability and conspecific densities: consequences for population regulation and range expansion

The presence of conspecifics can strongly modulate the quality of a breeding site. Both positive and negative effects of conspecifics can act on the same individuals, with the final balance between its costs and benefits depending on individual characteristics. A particular case of inter-individual variation found in many avian species is chromatic variability. Among birds, plumage coloration can co-vary with morphology, physiology and behavior as well as with age. These relationships suggest that cost-benefit balances of conspecific presence may be different for individuals with different colorations. We investigated whether inter-individual variability affects population regulation and expansion processes by analyzing potential differences in density-dependent productivity and settlement patterns in relation to plumage coloration in a population of a long-lived avian species recently undergoing a notable increase in numbers and distribution range. Our results show strong variation in the effect of density on productivity of breeding pairs depending on plumage coloration of their members. Productivity of dark birds decreased along the breeding density gradient while that of lighter breeders remained unchanged with conspecific density. In a similar way, our results showed an uneven occupation of localities by individuals with different plumage coloration in relation to local densities, with the breeding of lighter harriers more aggregated than that of dark-brown ones. At a population scale, darker birds had higher probability of colonization of the most isolated, empty sites. Explanations for species range expansion and population regulation usually make the inferred assumption that species traits are similar among individuals. However, in most species, there could be individual variation in niche requirements or dispersal propensities among individuals with different traits. Our results contribute to the growing appreciation that the individual traits, but not the average trait at the level of species, are important during population regulation and expansion processes.     

Acoustic Competition in Male Midwife Toads Alytes obstetricans and Alytes cisternasii: Response to Neighbor Size and Calling Rate. Implications for Female Choice

Effects of neighbor on male calling behavior was studied through playback experiments of synthetic calls to males of two species of midwife toads. The responses of resident males were scored considering two temporal parameters (call duration and calling rate) and one spectral parameter (dominant frequency). The sounds used for the playback tests included two levels of fundamental frequency (correlated with male size) and two levels of call repetition rate. In both species, resident males only changed their calling rate in the presence of an intruder, and the response was different for synthetic calls with two levels of dominant frequencies and with two calling rates. Resident size was not significantly correlated with the magnitude of the change in the calling rate. On the other hand, resident calling rate was significantly and positively correlated with the magnitude of the increase in calling rate of the stimulus. The maximum relative increase in calling rate was observed in A. cisternasii. In phonotaxis tests, females are preferentially attracted to calls emitted at a higher rate confirming the importance of changes in calling rate for female attraction.