Inference about density and temporary emigration in unmarked populations

Few species are distributed uniformly in space, and populations of mobile organisms are rarely closed with respect to movement, yet many models of density rely upon these assumptions. We present a hierarchical model allowing inference about the density of unmarked populations subject to temporary emigration and imperfect detection. The model can be fit to data collected using a variety of standard survey methods such as repeated point counts in which removal sampling, double-observer sampling, or distance sampling is used during each count. Simulation studies demonstrated that parameter estimators are unbiased when temporary emigration is either "completely random" or is determined by the size and location of home ranges relative to survey points. We also applied the model to repeated removal sampling data collected on Chestnut-sided Warblers (Dendroica pensylvancia) in the White Mountain National Forest, U.S.A. The density estimate from our model, 1.09 birds/ha, was similar to an estimate of 1.11 birds/ha produced by an intensive spot-mapping effort. Our model is also applicable when processes other than temporary emigration affect the probability of being available for detection, such as in studies using cue counts. Functions to implement the model have been added to the R package unmarked.     

Overwinter Survival of Neotropical Migratory Birds in Early-Successional and Mature Tropical Forests

Many Neotropical migratory species inhabit both mature and earty-successional forests on their wintering grounds, yet comparisons of survival rates between habitats are lacking. Consequently, the factors affecting habitat suitability for Neotropical migrants and the potential effects of tropical deforestation on migrants are not well understood. We estimated overwinter survival and capture probabilities of Wood Thrush ( Hylochichla mustelina ), Ovenbird ( Seiurus aurocapillus ), Hooded Warbler ( Wilsonia citrina ), and Kentucky Warbler ( Oporornis formosus ) inhabiting two common tropical habitat types, mature and early-successional forest. Our results suggest that large differences (for example, ratio of survival rates (γ) ≤ 0.85) in overwinter survival between these habitats do not exist for any of these species. Age ratios did not differ between habitats, but males were more common in forest habitats and females more common in successional habitats for Hooded Warblers and Kentucky Warblers. Future research on overwinter survival should address the need for age- and sex-specific survival estimates before we can draw strong conclusions regarding winter habitat suitability. Our estimates of overwinter survival extrapolated to annual survival rates that were generally lower than previous estimates of annual survival of migratory birds. Capture probability differed between habitats for Kentucky Warblers, but our results provide strong evidence against large differences in capture probability between habitats for Wood Thrush, Hooded Warblers, and Ovenbirds. We found no temporal or among-site differences in survival or capture probability for any of the four species. Additional research is needed to examine the effects of winter habitat use on survival during migration and between-winter survival.     

Cover Caption

Cover: Radar imagery can be used to map the locations from which migratory birds depart and to highlight important stopover sites. Shown are movements of long-distance migrants, such as the Blackburnian Warbler ( Dendroica fusca ), detected during spring migration near Buffalo, New York. Warbler photo by Charles Eiseman. See pages 440–448.     

Effects of Selection Cutting and Landscape-Scale Harvesting on the Reproductive Success of Two Neotropical Migrant Bird Species

Abstract: We measured the effects of forest harvesting at two spatial scales on the productivity of two Neotropical migrant bird species, the Black-throated Blue Warbler (   Dendroica caerulescens ) and the Ovenbird (  Seiurus aurocapillus ). We selected two forest landscapes representing extremes in harvesting intensity for northwestern New Brunswick, Canada. Within each landscape, we selected study plots located either in recent (<5 years) selection-cut stands (30% removal every 20 years) or in stands that were lightly harvested at least 30 years ago. Black-throated Blue Warblers reached higher densities in selection cuts, but their reproductive performance ( pairing and fledging success) did not differ significantly between stand types (uncut vs. selection cut) or between landscape types (intensively vs. moderately harvested). In 1998, nonetheless, 61% of the estimated number of fledglings produced in our plots were born in selection cuts. In contrast, this proportion was only 23% for Ovenbirds in 1998 and 1999 combined. Ovenbirds had lower densities, and their reproductive performance was significantly lower in selection cuts than in uncut plots. These results indicate that the effects of selection cutting on demography are species-specific and that Ovenbird persistence in selection cuts may be compromised unless the intensity or frequency of cutting is decreased.     

The role of distance,familiarity, and time of day in Carolina wrens responses to conspecific songs

Summary We tested two predictions of the ranging hypothesis (Morton 1982) which suggests that: (1) For distance estimation of singing intruders from them, birds compare degraded songs they hear to undegraded songs in their memory. (2) The strength of response of birds to songs depends on how well they can range the songs as coming from inside or outside the territory.In accordance with first prediction we found that, Carolina wrens discriminate significantly between songs played from outside to those played inside their territories only if they are familliar with the song type played (Fig. 1). In accordance with the second prediction, on the average, the birds in our study responded stronger to unfamiliar than to familiar songs when these were played from outside the territory, but stronger to familiar than to unfamiliar songs when played inside it (Fig. 2). We concluded that this was because they could range the familiar songs with greater certainty.The variance in the data was, in part, due to changes in the birds' responses during the day. We report for the first time, an increase in the strength of territorial defense of birds later in the morning, accompanied by a decrease in singing in response to song playback. We suggest that these changes reflect an increase in energy gain and worse acoustic conditions later in the day.     

Song repertoires do not hinder neighbor-stranger discrimination

Summary Through playback experiments and a comparative study we tested the hypothesis that song repertoires hinder individual recognition by song in birds. Our playback tests take advantage of the peculiar singing behavior of two species of parulid warblers: the American redstart (Setophaga ruticilla) and the yellow warbler (Dendroica petechia). Males of both species sing either the same song repeatedly (repeat mode) or a repertoire of songs in series (serial mode). Subjects were played the repeat song and the serial songs from a territorial neighbor and from a stranger some territories distant. Counter to the prediction of the hypothesis, they discriminated equally well between neighbors and strangers when the several serial songs were used as when the single repeat song was played. Thus, neither the small repertoires used by redstarts in serial mode (2–8 songs) nor the larger ones used by yellows (an average of over 20 songs) appear to hinder neighbor-stranger discrimination. In our comparative test, we examined the relationship between neighbor-stranger discrimination and repertoire size among 20 species for which data were available. Using two comparative methods, we did not find the predicted significant negative relationship.     

Song ranging by the dusky antbird,Cercomacra tyrannina: ranging without song learning

In a population of dusky antbirds (Cercomacra tyrannina), less aggressive responses to distance-degraded playbacks than to undegraded playbacks of pair duets show that this tropical suboscine passerine uses sound degradation to range distance from singing conspecifics. This is the first example of song-ranging in a species that does not learn songs, supporting the hypothesis that ranging preceded the song learning that occurs in more recently evolved passerine birds (oscines). Both sexes sing and are able to use song degradation to range distance from singers when their sex-specific song is played back. Received: 27 May 1994/Accepted after revision: 24 March 1996     

Sensitivity analysis of North American bird population estimates

The Partners in Flight North American Landbird Conservation Plan provided estimates of population sizes for 448 landbird species using a multiplicative model. Input parameters in this calculation included the area of state × Bird Conservation Region polygons, area-specific mean Breeding Bird Survey counts circa 1995, and adjustment factors for the distance over which species may presumably be correctly counted, the assumed pairing of singing males with non-singing females, and variability in the propensity of birds to sing over the course of the survey day. I assessed the sensitivity of this population calculation to changes in the input parameters. I assessed both local and global sensitivity of the model to changes in the parameters with Monte Carlo one-at-a-time simulations and the Fourier amplitude sensitivity test (FAST). Monte Carlo simulations were an estimate of local model sensitivity whereas FAST estimated global model sensitivity, accommodating the potential shared variance between model parameters. Monte Carlo simulations suggested population estimates were 39% more sensitive to changes in the detection distance adjustment than to the other parameters; the other parameters were nearly equal in their contribution to model sensitivity. Conversely, FAST analysis determined that each of the input variables aside from the pair adjustment provided roughly equal contributions to variability in population estimates. The most efficient means for improving continental population estimates for birds surveyed by the Breeding Bird Survey will be through increased scrutiny of the species-specific distance detection and time-of-day adjustments and improved understanding in the spatial and temporal variability in the mean Breeding Bird Survey count.     

Impacts of Chronic Anthropogenic Noise from Energy‐Sector Activity on Abundance of Songbirds in the Boreal Forest

Abstract: The effects of human activities in forests are often examined in the context of habitat conversion. Changes in habitat structure and composition are also associated with increases in the activity of people with vehicles and equipment, which results in increases in anthropogenic noise. Anthropogenic noise may reduce habitat quality for many species, particularly those that rely on acoustic signals for communication. We compared the density and occupancy rate of forest passerines close to versus far from noise‐generating compressor stations and noiseless well pads in the boreal forest of Alberta, Canada. Using distance‐based sampling, we found that areas near noiseless energy facilities had a total passerine density 1.5 times higher than areas near noise‐producing energy sites. The White‐throated Sparrow (Zonotrichia albicollis), Yellow‐rumped Warbler (Dendroica coronata), and Red‐eyed Vireo (Vireo olivaceus) were less dense in noisy areas. We used repeat sampling to estimate occupancy rate for 23 additional species. Seven had lower conditional or unconditional occupancy rates near noise‐generating facilities. One‐third of the species examined showed patterns that supported the hypothesis that abundance is influenced by anthropogenic noise. An additional 4 species responded negatively to edge effects. To mitigate existing noise impacts on birds would require approximately $175 million. The merits of such an effort relative to other reclamation actions are discussed. Nevertheless, given the $100 billion energy‐sector investment planned for the boreal forest in the next 10 years, including noise suppression technology at the outset of construction, makes noise mitigation a cost‐effective best‐management practice that might help conserve high‐quality habitat for boreal birds.     

Ranging of songs with the song type on use of different cues in Carolina wrens: effects of familiarity

Male territorial song birds are usually spaced far apart and most often hear conspecific song after it has been degraded by propagation through the environment. Their ability to use the degradation of songs to assess the distance of a singing rival without approaching (called ranging) presumably increases the efficiency of defending a territory. In order to assess degradation in a song the receiver needs to compare the characteristics of the received song to its characteristics at the source or at different distances. Earlier experiments on ranging in species with song repertoires have suggested that prior familiarity with the particular song type is necessary for ranging. Here I show that male Carolina wrens (Thryothorus ludovicianus) can use either temporal or spectral characteristics for ranging song types which they were unlikely to have heard previously. Playbacks consisting of only one song prevented subjects' close-range experience with the loudspeaker, and flights beyond the loudspeaker provided direct evidence for over-assessment of distance when songs were degraded. Because ranging of songs was not affected by the degree of familiarity with the song type, this experiment provides no evidence that song repertoires hinder ranging in Carolina wrens, as suggested by Morton's ranging hypothesis. Instead, at least approximate ranging of songs is evidently possible by assessment of degradation in general features of a species' songs. Received: 9 August 1996 / Accepted after revision: 15 February 1997     

Long-Term Effects of Group-Selection Timber Harvesting on Abundance of Forest Birds

Abstract:  Relatively few studies have examined the ecological effects of group-selection timber harvesting, and nearly all have been short-term and have lacked experimental manipulations that allow pre- and posttreatment comparisons. We have been documenting the effects of a group-selection timber harvest on bird abundance in a Maine forest for 24 years (preharvest, 1983–1987; postharvest, 1988–2006). Here we characterized the trends in bird abundance over the first 20 years of the study in the managed and control halves of the 40-ha study area. Species responses to the group-selection harvest were idiosyncratic, but in general the mature-forest bird community was retained and species dependent on early successional habitat temporarily (≤8 years) benefited. The Eastern Wood-Pewee ( Contopus virens ) , Winter Wren ( Troglodytes troglodytes ) , Pine Warbler ( Dendroica pinus ) , and White-throated Sparrow ( Zonotrichia albicollis ) increased in abundance in the managed half of the study area following timber harvest, whereas the Veery ( Catharus fuscescens ) decreased. The Black-and-White Warbler ( Mniotilta varia ) , Nashville Warbler ( Vermivora ruficapilla ) , and Common Yellowthroat ( Geothlypis trichas ) responded positively to harvesting, as indicated by decreases in abundance in the control area and more protracted declines or stable abundances in the managed area. This study constitutes the longest experimental investigation to date of the effects of a group-selection harvest on birds and thus provides important information on the strength, direction, and duration of temporal changes in bird populations following forest management.     

Individual recognition of song repertoires in two wood warblers

We examined recognition of individual neighbors in hooded warblers (Wilsonia citrina) and Kentucky warblers (Oporornis formosus) by comparing responses to playbacks of neighbors' songs from appropriate and inappropriate boundaries of the subjects' territories. Kentucky warblers, which sing one song type each, responded more strongly to the songs of neighbors broadcast from incorrect than from correct boundaries. Hooded warblers use their repertoires of three to nine song types in two distinct modes of singing: repeat mode involves repeated presentation of one song type; mixed mode involves presentation of two or more song types in irregular sequence. Playbacks of neighbors' mixed-mode songs from appropriate and inappropriate boundaries indicated capabilities of individual recognition similar to those reported previously for repeat-mode songs (Godard 1991) and to those of Kentucky warblers. Repertoires of moderate size, therefore, have no pronounced influence on eventual recognition of individual neighbors in hooded warblers.     

Sound degradation as a distance cue in great tit (Parus major) song

Summary We report an experiment designed to test the ideas that: 1. male songbirds can use cues from the distortion of song by environmental factors (degradation) to estimate the distance of another singing male; 2. song degradation is assessed by reference to an internal standard. Great tits respond more strongly to undegraded than to degraded songs when both are played at the same amplitude and from the same position in the territory. This difference in response is shown only if the playback song is familiar to the test bird; familiar songs being those sung either by the test bird or neighbours of the test bird. We interpret these results as evidence that cues from song degradation can be used to estimate the distance of a singing conspecific and that degradation assessment is only possible if the bird has an internal representation of the song (because either it and/or a neighbour sings the song). We discuss the implications of these results for Morton's (1982) ranging hypothesis, and for the distinction between learning and performance in bird song. Our results partially support the ranging hypothesis, but question the nature of unrangeable songs sensu Morton. The finding that birds can assess the degree of degradation of songs that they do not sing, supports the idea that birds learn more songs than they sing.     

Choice of singing sites by male bushcrickets (Tettigonia viridissima) in relation to signal propagation

Summary The spatial dispersion of singing male bushcrickets (Tettigonia viridissima) in a marshland habitat was found to be significantly clumped. Males clustered in patches of taller vegetation, buth within such clusters, males were regularly spaced with a mean distance of 6 m between nearest-neighbours. Males occupied perches on plants that were, on average, 0.3 m above the top of surrounding vegetation. Physical and acoustical interactions between males were observed more frequently when males were singing from higher sites. Excess attenuation of the male song was found to increase with frequency but decreased markedly with increasing elevation of the singing male above the ground. The maximum detection range of the song, realized when the insect was singing > 1 m above the surrounding vegetation, was estimated as 60 m for the fundamental frequency (10 kHz), 38 m for the 1st harmonic (20 kHz) and 14 m for the second harmonic (30 kHz). By contrast, when males sang from the middle of dense reed beds, the estimated detection distance was only 8 m, 6 m and 4 m for each frequency band, respectively. Males could have increased the detection range of their songs almost three fold by singing from higher positions than those usually observed in the field. This suggests that there may be a cost of singing at higher elevations such as an increased risk of predation and/or increased aggression from neighbours. We suggest the spacing strategy adopted by males reflects a compromise between maximizing the range over which their songs can be detected and accurately localized by females and minimizing interference from competing males. Offprint requests to: T. Eiriksson     

Use of fecundity measured directly throughout the breeding season to test a source-sink demographic model

Populations of landbirds (bird species that occupy terrestrial habitats for most of their life cycle) are declining throughout North America (north of Mexico) and Europe, yet little is known about how demography is driving this trend. A recent model of 5 geographically separated populations of Cerulean Warblers (Dendroica cerulea) that was based on within-season sampling of nest survival and fledgling success shows that all populations are sinks (annual reproduction is consistently less than annual adult mortality). I tested this indirect model by directly measuring fecundity (number of female fledglings/female) during the breeding season for 2 years in a Cerulean Warbler population occupying a mature forest in southwestern Michigan (U.S.A.) I determined territories of male birds on the basis of male plumage characters and phases of the nesting cycle (2007) and on uniquely color-banded males (2008). I transferred locations of identified males to topographic maps. I counted all fledglings in territories from May to July each year. The model I tested may apply only to single-brooded species; therefore, I searched the literature to estimate the percentage of single-brooded species in North America. The breeding season of Cerulean Warblers was short- nearly all nests were initiated from mid-May to late June. Nest predation and brood parasitism were primary and rare causes of nest failure, respectively. Significantly fewer Cerulean Warblers fledged from parasitized than from nonparasitized nests. Fledgling survival required to maintain the population size was well above previously published values for Neotropical migrants. Single-brooded species comprise 62% of North American breeding bird species for which the number of broods per year is known; I believe my results may apply to these species. The consistency between identification of populations as sources or sinks on the basis of either model estimates or direct measurements suggests that a demographic model relying on within-season sampling of fecundity is adequate to determine population status of single-brooded avian populations. In addition, on the basis of results of previous studies, annual adult survival rate of the Cerulean Warbler is typical of parulid warblers that are not declining. Thus, low fecundity, here determined with different quantitative methods, can drive status of landbird species with high-observed survival.     

Estimating the complexity of bird song by using capture-recapture approaches from community ecology

Repertoire size, the number of unique song or syllable types in the repertoire, is a widely used measure of song complexity in birds, but it is difficult to calculate this exactly in species with large repertoires. A new method of repertoire size estimation applies species richness estimation procedures from community ecology, but such capture-recapture approaches have not been much tested. Here, we establish standardized sampling schemes and estimation procedures using capture-recapture models for syllable repertoires from 18 bird species, and suggest how these may be used to tackle problems of repertoire estimation. Different models, with different assumptions regarding the heterogeneity of the use of syllable types, performed best for different species with different song organizations. For most species, models assuming heterogeneous probability of occurrence of syllables (so-called detection probability) were selected due to the presence of both rare and frequent syllables. Capture-recapture estimates of syllable repertoire size from our small sample did not differ significantly from previous estimates using larger samples of count data. However, the enumeration of syllables in 15 songs yielded significantly lower estimates than previous reports. Hence, heterogeneity in detection probability of syllables should be addressed when estimating repertoire size. This is neglected using simple enumeration procedures, but is taken into account when repertoire size is estimated by appropriate capture-recapture models adjusted for species-specific song organization characteristics. We suggest that such approaches, in combination with standardized sampling, should be applied in species with potentially large repertoire size. On the other hand, in species with small repertoire size and homogenous syllable usage, enumerations may be satisfactory. Although researchers often use repertoire size as a measure of song complexity, listeners to songs are unlikely to count entire repertoires and they may rely on other cues, such as syllable detection probability.Communicated by A. Cockburn     

Responses to playback of whistle songs and normal songs in male nightingales: effects of song category, whistle pitch, and distance

In most species of song birds, males develop song repertoires of several different songs. Among this variety, different songs may be used differently in communication and, thus, may have different functions. Here we studied vocal responses to playback of structurally different songs in male territorial nightingales (Luscinia megarhynchos). Nightingales have enormous vocal repertoires of about 200 song types. Among these songs, one category, whistle songs, sticks out syntactically and acoustically. Here we tested whether or not male nightingales match whistle songs with whistle songs and, if so, whether they also match the pitch of the broadcast whistles. Furthermore, we also tested if nightingales treat whistle songs as a separate category of songs. We conducted interactive playback experiments on nocturnal song in which each male received three playback treatments that differed in the number of whistle songs broadcast. Males responded differently during playbacks by singing significantly more whistle songs when the playback tapes contained many whistle songs than when they contained no whistle songs. Males also frequently matched the pitch of the broadcast whistle songs. In contrast to responses during playback, after the playback terminated males sang more whistle songs when no whistle songs were broadcast than when many whistle songs were broadcast. These findings suggest that whistle songs have a specific signal value and that nightingales treat them as a special song category. Electronic supplementary material to this paper can be obtained by using the Springer Link server located at http://dx.doi.org/10.1007/s00265-002-0511-1.     

Habituation and song repertoires in the great tit

Summary This paper is concerned with the idea that the song repertoires of passerine birds are an evolutionary adaptation to reduce habituation in listeners. In an experiment involving 16 territorial males I played either a single song or a repertoire of songs for 15 two-min trials through a single loudspeaker near the edge of the territory. In a second experiment with 10 birds I played the songs through one of two loudspeakers in different parts of the territory, alternating between loudspeakers on successive trials. The birds tended to habituate more rapidly to single song playback than to repertoires. In the second experiment the overall level of habituation was lower and the difference between the two treatments was less marked.Two features of song repertoire organisation are consistent with the habituation hypothesis (1) the avoidance of low recurrence intervals in switches between song types, and (2) the fact that within repertoire variability is as great or greater than between repertoire varability.The main problem with the habituation hypothesis is that habituation by listeners does not seem to be adaptive, so it is not clear why they should habituate. I suggest a hypothesis. Intruders may assess the density of birds in an area by listening to songs, so that habituation may be a mechanism by which this density assessment is achieved. Repertoires could be a mechanism by which resident birds cheat, through increasing the apparent density of singing birds.     

Voices in the dark: predation risk by owls influences dusk singing in a diurnal passerine

Predation is an important cost of communication in animals and thus a potent selection pressure on the evolution of signaling behavior. Heterospecific eavesdropping by predators may increase the vulnerability of vocalizing prey, particularly during low light, such as at dusk when nocturnal predators are actively hunting. Despite the risk it entails, dawn and dusk chorusing is common in passerines. However, the dusk chorus has not been studied much, neglecting the opportunity for understanding how eavesdropping between predators and prey may shape communication in birds. Here, we report the first demonstration of simulated predation risk (playback of owl vocalizations) altering the dusk chorus of a diurnal passerine, the veery (Catharus fuscescens). Veeries have a pronounced dusk chorus, singing well after sunset and potentially exposing themselves to predation by owls. In response to brief playbacks of owl calls (~30 s of calls presented three times over 25 min), veeries sang fewer songs post-sunset and stopped singing earlier relative to control trials. These changes in singing remained evident 30 min after the last owl stimulus. Although the avian dusk chorus has received relatively little attention to date, our results suggest that the dusk chorus may pose a higher predation risk to singing males that may influence the evolution of singing behavior in diurnal birds.     

Experimental investigation of false positive errors in auditory species occurrence surveys

False positive errors are a significant component of many ecological data sets, which in combination with false negative errors, can lead to severe biases in conclusions about ecological systems. We present results of a field experiment where observers recorded observations for known combinations of electronically broadcast calling anurans under conditions mimicking field surveys to determine species occurrence. Our objectives were to characterize false positive error probabilities for auditory methods based on a large number of observers, to determine if targeted instruction could be used to reduce false positive error rates, and to establish useful predictors of among-observer and among-species differences in error rates. We recruited 31 observers, ranging in abilities from novice to expert, who recorded detections for 12 species during 180 calling trials (66,960 total observations). All observers made multiple false positive errors, and on average 8.1% of recorded detections in the experiment were false positive errors. Additional instruction had only minor effects on error rates. After instruction, false positive error probabilities decreased by 16% for treatment individuals compared to controls with broad confidence interval overlap of 0 (95% CI:--46 to 30%). This coincided with an increase in false negative errors due to the treatment (26%;--3 to 61%). Differences among observers in false positive and in false negative error rates were best predicted by scores from an online test and a self-assessment of observer ability completed prior to the field experiment. In contrast, years of experience conducting call surveys was a weak predictor of error rates. False positive errors were also more common for species that were played more frequently but were not related to the dominant spectral frequency of the call. Our results corroborate other work that demonstrates false positives are a significant component of species occurrence data collected by auditory methods. Instructing observers to only report detections they are completely certain are correct is not sufficient to eliminate errors. As a result, analytical methods that account for false positive errors will be needed, and independent testing of observer ability is a useful predictor for among-observer variation in observation error rates.