Vocal Individuality of Great Gray Owls in the Sierra Nevada

ABSTRACT The cryptic plumage and nocturnal nature of the great gray owl (Strix nebulosa) make it difficult to study in its densely forested habitat. We investigated whether the vocalizations of individual great gray owls could be distinguished and used as a tool for population survey and monitoring. We recorded 312 territorial calls produced by 14 male and 11 female great gray owls between March and July 2006 and 2007 in the Sierra Nevada range of California, USA. We recorded 19 owls on multiple occasions within a season and 8 owls between seasons. We extracted 17 frequency and 15 temporal variables from the sonograms of each call. Discriminant analysis selected 9 variables and classified 92.8% of calls to the correct individual within a season; 71.4% of calls were classified to the correct individual between seasons. Our results indicate that territorial calls could be used to monitor individual great gray owls for both short- and long-term studies. Vocal individuality could be useful as a noninvasive method to improve census estimates and yield information on site fidelity, turnover rates, seasonal movements, and behavioral traits of great gray owls.     

Effects of episodic bamboo mast seeding on top predators in the southern Andes

Woody bamboos that undergo masting on a cyclic basis constitute large‐scale endogenous disturbances in forests of America, Africa and Asia, driving long‐ and short‐term effects on community structure and dynamics. Among the transient effects of these nonequilibrial phenomena are rodent outbreaks whose potential bottom‐up consequences on top predators have never been explored. We investigated the effects of unpredictable rodent outbreaks on the assemblage of nocturnal raptors of the southern Andes after a large‐scale (>140 000 ha), spatially heterogeneous, Chusquea culeou masting event in north Argentine Patagonia. We compared owl numbers and behaviours between pre‐masting (2009) and post‐masting (2011) at subsidized (outbreaking rodents) and unsubsidized (normal rodents) contiguous sites. Both generalist (opportunistic forest resident) and rodent‐specialist (forest‐facultative) owls were monitored, with emphasis on the resident territorial rufous‐legged owl (Strix rufipes). The resident owls behaved as predicted, perceiving the rodent increases soon and gathering at subsidized sites, while apparently relaxing territoriality. Contrary to our predictions, later at the rodent outbreak phase, resident territorial owls turned inconspicuous, coinciding – causally or not – with an irruption of forest‐facultative barn owls (Tyto alba tuidara), and influx of some open country short‐eared owls (Asio flammeus suinda, some of which took a chance to breed in the woods). Considering the whole rodent outbreak period, besides significant changes in owls’ numbers, we recorded a notable adjustment in owls’ foraging modes in response to food surplus (consuming prey heads only), and null interference behaviours among all observed species. This study provides a first quantitative assessment of the effects of bamboo episodic masting on top carnivores globally, and contributes novel data on the indirect effects of these events in forests of South America.     

Barred owl occupancy surveys within the range of the northern spotted owl

The range expansion by barred owls (Strix varia) into western North America has raised considerable concern regarding their potential effects on declining northern spotted owl (Strix occidentalis caurina) populations, yet most information on the occurrence of barred owls in the region is limited to incidental detections during surveys for spotted owls. To address this shortcoming we investigated response behavior, detection probabilities, and landscape occupancy patterns of barred owls in western Oregon, USA, during conspecific versus spotted owl call-broadcast surveys. Subtle differences in barred owl response behavior to conspecific versus spotted owl vocalizations combined with minor procedural differences between species-specific survey protocols led to a sizeable difference in estimated detection probabilities during conspecific (0.66, 95% CI = 0.61–0.71) versus spotted owl (0.48, 95% CI = 0.39–0.56) surveys. We identified 61 territorial pairs of barred owls during repeated surveys of a multi-ownership study area with the probability of occupancy being highest in the structurally diverse mixture of mature and old forests that occurred almost entirely on public lands. Our findings suggest that research and management strategies to address potential competitive interactions between spotted owls and barred owls will require carefully designed, species-specific survey methods that account for erratic response behaviors and imperfect detection of both species. Our sampling methods can be used by forest managers to determine the occurrence and distribution of barred owls with high confidence. © 2011 The Wildlife Society.     

Site Occupancy Dynamics of Northern Spotted Owls in the Eastern Cascades,Washington, USA, 1990–2003

Abstract: Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. Several spotted owl (Strix occidentalis) response variables have been examined in various investigations, but recent advances in statistical modeling permit evaluations of temporal and spatial variability in site occupancy, local-extinction, and colonization probabilities while incorporating imperfect detection probabilities. Following recent work by other researchers on site occupancy dynamics of spotted owls in Oregon, USA, we evaluated temporal variability of detection, occupancy, local-extinction, and colonization probabilities for spotted owls, as well as potential influences of barred owl (Strix varia) presence on these parameters. We used spotted owl survey data collected from 1990 to 2003 on a study area in the eastern Cascades Mountains, Washington, USA, to compare competing occupancy models from Program PRESENCE using Akaike's Information Criterion. Detection probabilities for individual spotted owls ranged from 0.54 to 0.80 if barred owls were not detected during the survey season and from 0.19 to 0.71 if barred owls were detected during the survey season. Pair detection probabilities ranged from 0.27 to 0.67 if barred owls were not detected during an individual survey and from 0.09 to 0.36 if barred owls were detected during an individual survey. During the study, site occupancy probabilities for spotted owl pairs declined by approximately 50%. For all spotted owls, both singles and pairs, site occupancy probabilities declined moderately during the study. Barred owl presence was negatively associated with spotted owl detection probabilities, and it had a positive association with local-extinction probabilities for all spotted owls, both singles and pairs. Given that our study area has supported higher densities of barred owls for longer periods than other study areas, our results may provide insight into how barred owls have influenced spotted owl site occupancy dynamics in adjacent British Columbia, Canada, or will influence spotted owl site occupancy dynamics in Oregon and California, USA, in the future.     

Factors affecting spontaneous vocal activity of Tawny Owls Strix aluco and implications for surveying large areas

To use vocalizations properly for the estimation of owl population size, it is important to identify how environmental factors affect owl calling behaviour. Here, we analyse how intrinsic and extrinsic factors modify the vocal activity of Tawny Owls Strix aluco in two areas of northern Spain. From March 2013 to May 2015, we radiotracked 20 Tawny Owls and also undertook a systematic survey in which we collected data on spontaneous vocal activity (hoot/call) of the tagged owls, plus their mates and neighbours (36 untagged owls). After 223 nights in Valle de Mena and 224 in Duranguesado we obtained a total of 8791 records of vocal activity. The annual proportion of surveys on which an owl called was 6.3% and did not differ between the study areas. Vocal activity of Tawny Owls varied with sex, annual cycle stage and weather. Male owls were significantly more vocal than females year‐round, and vocal activity peaked during the incubation and post‐breeding periods. Wind and rain adversely affected vocal activity of both sexes throughout the year. Tagged owls were detected more often than untagged owls, which we interpret as an observer effect because the systematic survey ensured that short distances to tagged owls increased the probability of detecting vocal activity. In fact, 2.8% of variation in vocal activity was due to detectability differences between tagged and untagged owls. We conclude that if fieldwork is carried out during the optimum period of the year for vocal detection (i.e. incubation, which in our case was around mid‐April), and under good weather conditions (dry and calm nights), censuses based on spontaneous vocal activity would detect only approximately 12% of the true Tawny Owl population.     

A case study of male tawny owl (Strix aluco) vocalizations in South Korea: call feature,individuality, and the potential use for census

Vocal individuality has been used as a monitoring tool, and two criteria are a prerequisite: high variation among individuals and low variation within individuals, and vocal consistency within and across seasons. We examined individual variation in the territorial hoot calls of the tawny owl (Strix aluco) to discriminate between males and to assess a possible conservation technique that would allow for monitoring individuals within a study area. The territorial calls were recorded from five males in the Naejang Mountain National Park in South Korea during the breeding season in 2015 and 2016 and analyzed both quantitatively and qualitatively to determine the amount of variation within and between individuals. Our results showed that the territorial calls were specific to individuals within a population and that the acoustic distances between males living in the same territory during the two years were the smallest for the four nesting sites. Our results suggest that territorial calls of the tawny owls are individually identifiable over two years and that this acoustic technique can be useful for monitoring individual site fidelity.     

Review of the effects of barred owls on spotted owls

Barred owls (Strix varia) are forest-dwelling owls, native to eastern North America, with populations that expanded westward into the range of the spotted owl (Strix occidentalis). Barred owls exert an overwhelmingly negative influence on spotted owls, thereby threatening spotted owl population viability where the species co-occur. In this review, we provide an overview of the barred owl's range expansion and detail and synthesize previously published literature on spotted and barred owls within the range of the spotted owl as related to potential future outcomes for the northern spotted owl (S. o. caurina). We include research on diet, habitat use and selection, effects of barred owls on spotted owl demography and behavior, hybridization with spotted owls, parasites, contemporary management, and future research needs for spotted owl populations given continued barred owl expansion throughout western North America. Our literature review and synthesis should provide managers with the information necessary to develop strategies that mitigate deleterious effects of barred owls at local and landscape scales. © 2019 The Wildlife Society.     

Variation in the male territorial hoot of the Tawny Owl Strix aluco in three English populations

Little is known about owl song. We made sonagrams of the territorial calls of 50 male Tawny Owls Strix aluco from three different areas. Six temporal and four frequency measures of the calls were recorded from the sonagrams. The measures of the calls were then subjected to analysis to try to separate between individual owls and between owls from different areas. We also looked for similarities between calls of neighbouring owls and for any effect of habitat on owl hoots. Individual owls were separated on the basis of their hoots with a high degree of success (98.6% overall), and there were significant differences between areas. Differences were found between calls in woodland and farmland habitats, but these differences were not in the direction expected to increase sound transmission. Calls of neighbouring owls did not resemble each other more than calls from owls that were not in vocal contact, implying that if calls are learned by Tawny Owls, they are learned before dispersal.     

Barred Owl Space Use and Habitat Selection in the Eastern Cascades,Washington

ABSTRACT Competition with barred owls (Strix varia varia) is an important factor contributing to the continued decline of threatened northern spotted owl (Strix occidentalis caurina) populations in the Pacific Northwest, USA, but basic information on habitat selection and space use patterns of barred owls is lacking for much of the region. We investigated space use and habitat selection by tracking radiotagged barred owls in the Eastern Cascade Range of Washington, USA, from 2004 to 2006. We surveyed for barred owls across the 309-km² study area and confirmed presence of barred owl pairs at 21 sites. We collected movement data on 14 barred owls from 12 sites. Mean annual 95% fixed-kernel home-range size was 194 ha for females (n = 4, SD = 70) and 288 ha for males (n = 5, SD = 114). Home ranges were located more frequently than expected in areas with low topographic position, gentle slopes, large overstory tree-crown diameter, high normalized difference vegetation index (NDVI), overstory tree canopy closure >72%, and a moderate amount of solar insolation. Within home ranges, areas that had large tree-crown diameters, low topographic positions, and gentle slopes were used more frequently than expected. The resource selection function we developed for barred owls in our study area indicated that barred owls used areas with the combination of low values for topographic position and slope and higher values for NDVI, solar insolation, and an interaction term for canopy closure and tree-crown diameter. In comparison to published information on northern spotted owls, barred owls used areas with similar canopy closure and tree size classes, but barred owl home ranges were much smaller and more concentrated on gentler slopes in valley bottoms. This information may contribute to the development of management practices that maintain forest characteristics appropriate for spotted owl habitat and prey in areas where spotted owls are least likely to be excluded by territorial barred owls in the Eastern Cascades of Washington.     

Microsatellite loci for distinguishing spotted owls (Strix occidentalis), barred owls (Strix varia), and their hybrids

We identified four diagnostic microsatellite loci that distinguish spotted owls (Strix occidentalis), barred owls (Strix varia), F₁ hybrids and backcrosses. Thirty‐four out of 52 loci tested (65.4%) successfully amplified, and four of these loci (11.8%) had allele sizes that did not overlap between spotted and barred owls. The probability of correctly identifying a backcross with these four loci is 0.875. Genotyping potential hybrid owls with these markers revealed that field identifications were often wrong. Given the difficulty of identifying hybrids in the field, these markers will be useful for hybrid identification, law enforcement and spotted owl conservation.     

Reproduction of northern spotted owls: The role of local weather and regional climate

We examined associations between annual reproduction and climate for 6 populations of individually marked northern spotted owls (Strix occidentalis caurina) in Washington and Oregon. We used an information-theoretical approach and mixed models to evaluate statistical models representing a priori hypotheses about the effects of weather and climate on reproduction. Reproduction was higher for adult than subadult owls and declined as the proportion of spotted owl territories with barred owl (Strix varia) detections increased. Similar to other spotted owl studies, we found that reproduction was negatively associated with cold, wet winters and nesting seasons at 3 of 6 study areas. In addition, we identified new relationships between reproduction, annual precipitation, storms, and regional climate cycles. For 3 of 6 areas, we found a quadratic relation between precipitation (rain and snow) and reproduction, with the number of young fledged per pair per year declining as precipitation in the previous year deviated from average levels. A meta-analysis conducted across all 6 areas indicated that reproduction at the regional level had a quadratic association with total winter snowfall in the preceding winter and was positively related to temperatures during the previous summer and fall. The amount of annual variation in reproduction accounted for by weather and climate varied widely across the 6 areas (4–79%), whereas variation in weather and climate across owl territories accounted for little of the spatial variation in reproduction (0–4%). Our results suggest that across the range of the species climate factors affecting prey abundance may have a greater effect on reproduction than direct effects of weather on nestlings. © 2011 The Wildlife Society.     

Interspecific offspring killing in owls

In Baranya County (Southern Hungary), tawny owls (Strix aluco) and barn owls (Tyto alba) sequentially use the same nest boxes in a significant number of cases. A total of 460 broods were observed between 1996 and 2003 and, in 12 cases, whole broods of dead tawny owl chicks were registered that had apparently been killed. On investigating the reproductive life characteristics, population sizes, and frequency of killing in these two species, it was concluded that: (1) with growing barn owl population, the number of sequential broods increases but changes in tawny owl population size have no effect on the frequency of sequential broods; (2) the number of killings depends on the number of sequential broods; and (3) with growing barn owl population, the number of killings also grows and this change is unaffected by the size of the tawny owl population. However, no killing occurs as long as 50–60% of the nest boxes are unoccupied. There is no killing either until the percentage of nest boxes occupied by barn owls reaches 40%, although a threshold value like this cannot be shown for the tawny owl. In the cases when a barn owl breeding follows the tawny owl's, the percentage of killing is significantly higher compared to that when barn owls do not breed in the same box. These results indicate that barn owls kill the offspring of tawny owls. By these means, they obtain a breeding place earlier than without killing the chicks of the other species, and this results in higher reproductive success. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 488–494.     

BIOACOUSTIC PUBLICATIONS, 1989

ABSTRACT

Coruros Spalacopus cyanus, social fossorial rodents from Chile, use a complex acoustic repertoire with eleven different true vocalisations and one mechanical sound in various behavioural contexts. The complex of contact calls is particularly well differentiated. Juvenile coruros produced six true vocalisations of which four were structurally identical to adult calls. One vocalisation had components of two adult sounds and one occurred only in juvenile animals. Certain calls from the adult repertoire were lacking. The frequencies of sounds of juveniles were considerably higher than those of adults, with many sounds reaching the ultrasonic range. Nevertheless, pure ultrasonic sounds were not recorded.

The frequencies of the analysed sounds of coruros extended from 0.17 to 20.33 kHz with dominant frequency components between 0.17 and 10 kHz. The acoustic properties of calls are suitable for transmission above and below ground, thus providing further indirect evidence that coruros are not strictly confined to an underground way of life. Indeed, the great variability of frequency ranges, with lower frequencies always being included, reflects a specialisation for communication in variable acoustic environments.

The most distinctive and unique vocalisation of coruros is the long duration musical trilling (lasting up to two minutes), which is a long-distance call emitted in alarm and arousal contexts. Recordings of this call from natural burrows in the field in Chile showed similar structural features to vocalisations from captive colonies in the laboratory.

Our findings provide a further example of matching physical properties of vocalisations to the acoustic conditions of the habitat. However, vocalisations in subterranean rodents consist almost exclusively of short-distance calls, the trilling of coruros being the notable exception. Since the selective pressure of the acoustic environment upon the evolution of short-distance vocalisations is probably minimal, we suggest that during their evolution, subterranean mammals have matched their vocalisations primarily to their hearing range and not directly to the acoustics underground. Hearing probably has been the primary target of natural selection, serving not only for communication but also for detection of predators (and, in carnivores, of prey).     

Site occupancy dynamics of northern spotted owls in managed interior Douglas fir forests,California, USA, 1995–2009

Northern spotted owls (Strix occidentalis caurina) have received intense research and management interest since their listing as a threatened species by the United States Fish and Wildlife Service in 1990. For example, public and private forest managers in the Pacific Northwest, USA, conduct surveys to determine presence or absence of spotted owls prior to timber harvest operations. However, although recently developed statistical methods have been applied to presence–absence data collected during research surveys, the effectiveness of operational surveys for detecting spotted owls and evaluating site occupancy dynamics is not known. We used spotted owl survey data collected from 1995 to 2009 on a study area in interior northern California, USA, to evaluate competing occupancy models from Program PRESENCE using Akaike's Information Criterion (AIC). During 1,282 individual surveys, we recorded 480 spotted owl detections (37.4%) and 13 barred owl (1.0%) detections. Average per visit detection probability (85% CL) for single and paired spotted owls was 0.93 (0.90–0.96) for informed daytime, stand-based searches and 0.47 (0.43–0.51) for nighttime, station-based surveys (estimated from the best model); the average per visit detection probability from the null model was 0.67 (0.64–0.70). Average pair-only detection probabilities were 0.86 (0.81–0.90) for informed daytime, stand-based searches and 0.23 (0.18–0.29) for nighttime, station-based surveys; the average per visit detection probability from the null model was 0.63 (0.58–0.68). Site occupancy for any owl declined from 0.81 (0.59–0.93) in 1995 to 0.50 (0.39–0.60) in 2009; pair occupancy declined from 0.75 (0.56–0.87) to 0.46 (0.31–0.61). Our results suggest that a combination of 1 informed stand and 2 station-based operational surveys can support determinations of spotted owl site status (either a single or a pair) at desired levels of confidence. However, our information was collected in an area where barred owls were rarely detected. Surveys conducted in areas that support well-established barred owl populations are likely to be less effective for determining presence or absence of spotted owls and may require more surveys and/or different survey methods to determine site status with confidence. © 2012 The Wildlife Society.     

Multistate Models Reveal Long-Term Trends of Northern Spotted Owls in the Absence of a Novel Competitor

Quantifying spatial and temporal variability in population trends is a critical aspect of successful management of imperiled species. We evaluated territory occupancy dynamics of northern spotted owls (Strix occidentalis caurina), California, USA, 1990–2014. The study area possessed two unique aspects. First, timber management has occurred for over 100 years, resulting in dramatically different forest successional and structural conditions compared to other areas. Second, the barred owl (Strix varia), an exotic congener known to exert significant negative effects on spotted owls, has not colonized the study area. We used a Bayesian dynamic multistate model to evaluate if territory occupancy of reproductive spotted owls has declined as in other study areas. The state-space approach for dynamic multistate modeling imputes the number of territories for each nesting state and allows for the estimation of longer-term trends in occupied or reproductive territories from longitudinal studies. The multistate approach accounts for different detection probabilities by nesting state (to account for either inherent differences in detection or for the use of different survey methods for different occupancy states) and reduces bias in state assignment. Estimated linear trends in the number of reproductive territories suggested an average loss of approximately one half territory per year (-0.55, 90% CRI: -0.76, -0.33), in one management block and a loss of 0.15 per year (-0.15, 90% CRI: -0.24, -0.07), in another management block during the 25 year observation period. Estimated trends in the third management block were also negative, but substantial uncertainty existed in the estimate (-0.09, 90% CRI: -0.35, 0.17). Our results indicate that the number of territories occupied by northern spotted owl pairs remained relatively constant over a 25 year period (-0.07, 90% CRI: -0.20, 0.05; -0.01, 90% CRI: -0.19, 0.16; -0.16, 90% CRI: -0.40, 0.06). However, we cannot exclude small-to-moderate declines or increases in paired territory numbers due to uncertainty in our estimates. Collectively, we conclude spotted owl pair populations on this landscape managed for commercial timber production appear to be more stable and do not show sharp year-over-year declines seen in both managed and unmanaged landscapes with substantial barred owl colonization and persistence. Continued monitoring of reproductive territories can determine whether recent declines continue or whether trends reverse as they have on four previous occasions. Experimental investigations to evaluate changes to spotted owl occupancy dynamics when barred owl populations are reduced or removed entirely can confirm the generality of this conclusion.     

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

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

Subspecific relationships and genetic structure in the spotted owl

Hierarchical genetic structure was examined in the three geographically-defined subspecies of spotted owl (Strix occidentalis) to define relationships among subspecies and quantify variation within and among regional and local populations. Sequences (522 bp) from domains I and II of the mitochondrial control region were analyzed for 213 individuals from 30 local breeding areas. Results confirmed significant differences between northern spotted owls and the other traditional geographically defined subspecies but did not provide support for subspecific level differences between California and Mexican spotted owls. Divergence times among subspecies estimated with a 936 bp portion of the cytochrome b gene dated Northern and California/Mexican spotted owl divergence time to 115,000–125,000 years ago, whereas California/Mexican spotted owl divergence was estimated at 15,000 years ago. Nested clade analyses indicated an association between California spotted owl and Mexican spotted owl haplotypes, implying historical contact between the two groups. Results also identified a number of individuals geographically classified as northern spotted owls (S. o. caurina) that contained haplotypes identified as California spotted owls (S. o. caurina). Among all northern spotted owls sampled (n=131), 12.9% contained California spotted owl haplotypes. In the Klamath region, which is the contact zone between the two subspecies, 20.3% (n=59) of owls were classified as California spotted owls. The Klamath region is a zone of hybridization and speciation for many other taxa as well. Analyses of population structure indicated gene flow among regions within geographically defined subspecies although there was significant differentiation among northern and southern regions of Mexican spotted owls. Among all areas examined, genetic diversity was not significantly reduced except in California spotted owls where the southern region consists of one haplotype. Our results indicate a stable contact zone between northern and California spotted owls, maintaining distinct subspecific haplotypes within their traditional ranges. This supports recovery efforts based on the traditional subspecies designation for the northern spotted owl. Further, although little variation was found between California and Mexican spotted owls, we suggest they should be managed separately because of current isolation between groups.     

Numerical and functional response of predators to a long-term decline in mammalian prey at a semi-arid Neotropical site

Summary Occurrence and diet of ten carnivorous predators (four falconiforms, four owls, and two foxes), and population levels of their mammalian prey, were monitored over 45 months at a semi-arid site in north-central Chile. Early in this period, small mammals irrupted and then declined markedly to a density 7% of that at peak. All four falconiforms (Buteo polyosoma, Falco sparverius, Geranoaetus melanoleucus, Parabuteo unicinctus) and one owl (Tyto alba) responded numerically to the decline in mammalian prey by virtually abandoning the study site. The three other owls (Athene cunicularia, Bubo virginianus, Glaucidium nanum) and the two foxes (Pseudalopex culpaeus and P. griseus) remained. With few exceptions, throughout the study predators maintained species-specific preferences among small mammal species regardless of the absolute and proportional abundance of these prev. At no time did the two prey species most responsible for the irruption (the rodents Phyllotis darwini and Akodon olivaceus) occur in predators' diets out of proportion to their estimated relative abundance in the field. Predators were clearly unable to prevent the irruption from occurring. Given the absence of a clear functional response to the most irruptive species, predators seemed unlikely to have been responsible for the observed crash. At present, however, predators may be prolonging the crash and delaying the return of small-mammal populations to typical densities.     

Individuality,kin similarity and experimental playback of contact calls in cooperatively breeding riflemen

Riflemen/tītipounamu (Acanthisitta chloris) are kin-based cooperatively breeding birds, which appear able to recognise their relatives. Here, we investigate the potential for vocalisations to act as recognition cues in riflemen. We identified an appropriate contact call and recorded it at the nest from 19 adult riflemen. Measurements of call characteristics were individually repeatable. In addition, call similarity was significantly correlated with relatedness among all birds and among males. Thus, in principle, these contact calls contain sufficient information for individual recognition of familiar kin, and some assessment of relatedness between unfamiliar birds. To test whether riflemen responded differently to calls of kin, we broadcast calls of relatives and non-relatives as separate treatments in a playback experiment. Focal birds rarely responded aggressively or affiliatively, and their tendency to do so was unrelated to treatment. We conclude that zip calls are suitable kin recognition cues, but whether they are used as such remains unknown.