Bird song and anthropogenic noise: vocal constraints may explain why birds sing higher-frequency songs in cities

When animals live in cities, they have to adjust their behaviour and life histories to novel environments. Noise pollution puts a severe constraint on vocal communication by interfering with the detection of acoustic signals. Recent studies show that city birds sing higher-frequency songs than their conspecifics in non-urban habitats. This has been interpreted as an adaptation to counteract masking by traffic noise. However, this notion is debated, for the observed frequency shifts seem to be less efficient at mitigating noise than singing louder, and it has been suggested that city birds might use particularly high-frequency song elements because they can be produced at higher amplitudes. Here, we present the first phonetogram for a songbird, which shows that frequency and amplitude are strongly positively correlated in the common blackbird (Turdus merula), a successful urban colonizer. Moreover, city blackbirds preferentially sang higher-frequency elements that can be produced at higher intensities and, at the same time, happen to be less masked in low-frequency traffic noise.     

Sleepless in Town – Drivers of the Temporal Shift in Dawn Song in Urban European Blackbirds

Organisms living in urban environments are exposed to different environmental conditions compared to their rural conspecifics. Especially anthropogenic noise and artificial night light are closely linked to urbanization and pose new challenges to urban species. Songbirds are particularly affected by these factors, because they rely on the spread of acoustic information and adjust their behaviour to the rhythm of night and day, e.g. time their dawn song according to changing light intensities. Our aim was to clarify the specific contributions of artificial night light and traffic noise on the timing of dawn song of urban European Blackbirds (Turdus merula). We investigated the onset of blackbird dawn song along a steep urban gradient ranging from an urban forest to the city centre of Leipzig, Germany. This gradient of anthropogenic noise and artificial night light was reflected in the timing of dawn song. In the city centre, blackbirds started their dawn song up to 5 hours earlier compared to those in semi-natural habitats. We found traffic noise to be the driving factor of the shift of dawn song into true night, although it was not completely separable from the effects of ambient night light. We additionally included meteorological conditions into the analysis and found an effect on the song onset. Cloudy and cold weather delayed the onset, but cloud cover was assumed to reflect night light emissions, thus, amplified sky luminance and increased the effect of artificial night light. Beside these temporal effects, we also found differences in the spatial autocorrelation of dawn song onset showing a much higher variability in noisy city areas than in rural parks and forests. These findings indicate that urban hazards such as ambient noise and light pollution show a manifold interference with naturally evolved cycles and have significant effects on the activity patterns of urban blackbirds.     

Clocks for the city: circadian differences between forest and city songbirds

To keep pace with progressing urbanization organisms must cope with extensive habitat change. Anthropogenic light and noise have modified differences between day and night, and may thereby interfere with circadian clocks. Urbanized species, such as birds, are known to advance their activity to early morning and night hours. We hypothesized that such modified activity patterns are reflected by properties of the endogenous circadian clock. Using automatic radio-telemetry, we tested this idea by comparing activity patterns of free-living forest and city European blackbirds (Turdus merula). We then recaptured the same individuals and recorded their activity under constant conditions. City birds started their activity earlier and had faster but less robust circadian oscillation of locomotor activity than forest conspecifics. Circadian period length predicted start of activity in the field, and this relationship was mainly explained by fast-paced and early-rising city birds. Although based on only two populations, our findings point to links between city life, chronotype and circadian phenotype in songbirds, and potentially in other organisms that colonize urban habitats, and highlight that urban environments can significantly modify biologically important rhythms in wild organisms.     

Experimental evidence for an impact of anthropogenic noise on dawn chorus timing in urban birds

Many animal species are living in urban areas, where they encounter human‐altered environmental conditions. Artificial light and traffic noise are two of the most prominent anthropogenic factors, both of which potentially affect animal life. Here we studied the changes in traffic noise conditions over the morning in the urban bird habitat of the city of Seville, Spain. We tested experimentally whether noise from human activities can cause a shift in the timing of birdsong activity. Our data revealed that noise conditions vary markedly among our replicate set of twelve streets. Relatively quiet streets show low base‐line amplitude levels early in the morning, with frequent events of brief noise bursts, followed by a strong rise in noise levels. Relatively noisy streets have high base‐line amplitude levels from a much earlier start in the morning. Experimental exposure data revealed a noise‐related earlier start of dawn singing for two out of six species: the spotless starling Sturnus unicolor and the house sparrow Passer domesticus. Our experiment did not cover earlier singing species and revealed no impact for species with more‐variable starting times of the dawn singing. Our study provides more insight into the intertwinings of bird and human behavior and confirms the potential for experimental approaches to successfully tackle questions related to the impact of anthropogenic factors on animal life in cities.     

Birdsong and anthropogenic noise: implications and applications for conservation

The dramatic increase in human activities all over the world has caused, on an evolutionary time scale, a sudden rise in especially low-pitched noise levels. Ambient noise may be detrimental to birds through direct stress, masking of predator arrival or associated alarm calls, and by interference of acoustic signals in general. Two of the most important functions of avian acoustic signals are territory defence and mate attraction. Both of these functions are hampered when signal efficiency is reduced through rising noise levels, resulting in direct negative fitness consequences. Many bird species are less abundant near highways and studies are becoming available on reduced reproductive success in noisy territories. Urbanization typically leads to homogenization of bird communities over large geographical ranges. We review current evidence for whether and how anthropogenic noise plays a role in these patterns of decline in diversity and density. We also provide details of a case study on great tits (Parus major), a successful urban species. Great tits show features that other species may lack and make them unsuitable for city life. We hypothesize that behavioural plasticity in singing behaviour may allow species more time to adapt to human-altered environments and we address the potential for microevolutionary changes and urban speciation in European blackbirds (Turdus merula). We conclude by providing an overview of mitigating measures available to abate noise levels that are degrading bird breeding areas. Bird conservationists probably gain most by realizing that birds and humans often benefit from the same or only slightly modified measures.     

Does light pollution alter daylength? A test using light loggers on free-ranging European blackbirds (Turdus merula)

Artificial light at night is one of the most apparent environmental changes accompanying anthropogenic habitat change. The global increase in light pollution poses new challenges to wild species, but we still have limited understanding of the temporal and spatial pattern of exposure to light at night. In particular, it has been suggested by several studies that animals exposed to light pollution, such as songbirds, perceive a longer daylength compared with conspecifics living in natural darker areas, but direct tests of such a hypothesis are still lacking. Here, we use a combination of light loggers deployed on individual European blackbirds, as well as automated radio-telemetry, to examine whether urban birds are exposed to a longer daylength than forest counterparts. We first used activity data from forest birds to determine the level of light intensity which defines the onset and offset of daily activity in rural areas. We then used this value as threshold to calculate the subjective perceived daylength of both forest and urban blackbirds. In March, when reproductive growth occurs, urban birds were exposed on average to a 49-min longer subjective perceived daylength than forest ones, which corresponds to a 19-day difference in photoperiod at this time of the year. In the field, urban blackbirds reached reproductive maturity 19 day earlier than rural birds, suggesting that light pollution could be responsible of most of the variation in reproductive timing found between urban and rural dwellers. We conclude that light at night is the most relevant change in ambient light affecting biological rhythms in avian urban-dwellers, most likely via a modification of the perceived photoperiod.     

Underlying physiological control of reproduction in urban and forest-dwelling European blackbirds Turdus merula

The development and the continual expansion of urban areas have not only destroyed natural habitats, but also have drastically changed the environmental and ecological conditions of these areas. Consequently, species that have settled in these new man-made ecosystems are exposed to considerable alternations in environmental conditions compared to their 'wild' conspecifics. To understand the impact of human-induced environmental changes on life history events such as reproduction, we compared the timing of the reproductive season and its underlying endocrine control in free-living European blackbirds Turdus merula inhabiting urban and nearby forest areas. Body mass, fat score, gonadal size, luteinizing hormone (LH), testosterone (T), and estradiol (E₂) were measured. Urban blackbirds developed their gonads approximately three weeks earlier than forest birds, whereas the timing of gonadal regression did not differ. There are several factors (e.g. artificial light, temperature, food availability, and social cues) which may have caused the differences in the temporal organization of gonadal growth between the urban and forest-living populations. The advanced gonadal development of urban blackbirds did not coincide with an earlier secretion of reproductive hormones. In contrast, urban males had lower plasma LH and T levels during testicular growth than forest males. Differences in social interactions and environmental conditions may explain the contrast of gonadal development and the timing of hormone secretion between urban and forest blackbirds.     

Maternal effects and urbanization: Variation of yolk androgens and immunoglobulin in city and forest blackbirds

Wildlife inhabiting urban environments exhibit drastic changes in morphology, physiology, and behavior. It has often been argued that these phenotypic responses could be the result of micro‐evolutionary changes following the urbanization process. However, other mechanisms such as phenotypic plasticity, maternal effects, and developmental plasticity could be involved as well. To address maternal effects as potential mechanisms, we compared maternal hormone and antibody concentrations in eggs between city and forest populations of European blackbirds (Turdus merula), a widely distributed species for which previous research demonstrated differences in behavioral and physiological traits. We measured egg and yolk mass, yolk concentrations of androgens (androstenedione [A₄], testosterone [T], 5α‐dihydrotestosterone [5α‐DHT], and immunoglobulins [IgY]) and related them to population, clutch size, laying order, embryo sex, and progress of breeding season. We show (a) earlier onset of laying in the city than forest population, but similar egg and clutch size; (b) higher overall yolk androgen concentrations in the forest than the city population (sex‐dependent for T); (c) greater among‐female variation of yolk T and 5α‐DHT concentrations in the forest than city population, but similar within‐clutch variation; (d) similar IgY concentrations with a seasonal decline in both populations; and (e) population‐specific positive (city) or negative (forest) association of yolk A₄ and T with IgY concentrations. Our results are consistent with the hypotheses that hormone‐mediated maternal effects contribute to differences in behavioral and physiological traits between city and forest individuals and that yolk androgen and immunoglobulin levels can exhibit population‐specific relationships rather than trade‐off against each other.     

Seasonal and urban effects on the endocrinology of a wild passerine

In order to maximize their fitness, organisms in seasonal environments rely on external cues to optimally time their life‐history stages. One of the most important zeitgeber to time reproduction is the photoperiod, but further environmental cues are assessed to fine‐tune reproduction due to year‐to‐year variation in environmental conditions. However, in urbanized environments, the pervasive artificial light at night has altered the natural signal of light and darkness. Accordingly, artificial light at night was repeatedly shown to affect avian reproductive physiology and to advance seasonal reproduction in birds. However, these experiments were mainly conducted in the absence of further environmental cues to facilitate the investigation of the mechanisms which are still poorly understood. Here, we investigate whether the endocrine system of free‐ranging European blackbirds (Turdus merula) correlates with the amount of artificial light at night along a rural to urban gradient while the birds still encounter complementary environmental cues including seasonal variation in day length and temperature. Testosterone and estrone were assessed as metabolites in fecal samples and corticosterone in blood from mist‐netted blackbirds. We demonstrate that seasonal fluctuations in abiotic factors, individual conditions, but also light at night affect the reproductive and stress physiology of wild European blackbirds. Elevated artificial night light intensities were significantly positively correlated with corticosterone and negatively with female estrone levels. No effects of artificial light were found for testosterone levels. Our results suggest that female blackbirds in particular perceive even low levels of artificial light at night as a weak but chronic stressor that interacts with the hypothalamic‐pituitary‐gonadal axis and leads to a reduced secretion of reproductive hormones. These findings point out that the impacts of light pollution are diverse and we only slowly disentangle its multiple effects on physiology, ecology, and biodiversity.     

Sex and seasonal differences in neurogenesis and volume of the song‐control system are associated with song in brood‐parasitic and non‐brood‐parasitic icterid songbirds

The song‐control system in the brain of songbirds is important for the production and acquisition of song and exhibits both remarkable seasonal plasticity and some of the largest neural sex differences observed in vertebrates. We measured sex and seasonal differences in two nuclei of the song‐control system of brood‐parasitic brown‐headed cowbirds (Molothrus ater) and closely‐related non‐parasitic red‐winged blackbirds (Agelaius phoeniceus). These species differ in both the development and function of song. Brown‐headed cowbirds have a larger sex difference in song than red‐winged blackbirds. Female cowbirds never sing, whereas female blackbirds do though much less than males. In cowbirds, song primarily functions in mate choice and males modify their song as they approach sexual maturity and interact with females. In red‐winged blackbirds, song is used primarily in territorial defence and is crystalized earlier in life. We found that the HVC was more likely to be discernable in breeding female blackbirds than in breeding female cowbirds. Compared to males, females had a smaller HVC and a smaller robust nucleus of the arcopallium (RA). However, females had higher doublecortin immunoreactivity (DCX+) in HVC, a measure of neurogenesis. Consistent with sex differences in song, the sex difference in RA volume was greater in cowbirds than in blackbirds. Males of both species had a smaller HVC with higher DCX+ in post‐breeding condition than in breeding condition when song is more plastic. Sex and seasonal differences in the song‐control system were closely related to variation in song in these two icterid songbirds. © 2016 Wiley Periodicals, Inc. Develop Neurobiol 76: 1226–1240, 2016     

Future Climate Change Will Favour Non-Specialist Mammals in the (Sub)Arctics

Arctic and subarctic (i.e., [sub]arctic) ecosystems are predicted to be particularly susceptible to climate change. The area of tundra is expected to decrease and temperate climates will extend further north, affecting species inhabiting northern environments. Consequently, species at high latitudes should be especially susceptible to climate change, likely experiencing significant range contractions. Contrary to these expectations, our modelling of species distributions suggests that predicted climate change up to 2080 will favour most mammals presently inhabiting (sub)arctic Europe. Assuming full dispersal ability, most species will benefit from climate change, except for a few cold-climate specialists. However, most resident species will contract their ranges if they are not able to track their climatic niches, but no species is predicted to go extinct. If climate would change far beyond current predictions, however, species might disappear. The reason for the relative stability of mammalian presence might be that arctic regions have experienced large climatic shifts in the past, filtering out sensitive and range-restricted taxa. We also provide evidence that for most (sub)arctic mammals it is not climate change per se that will threaten them, but possible constraints on their dispersal ability and changes in community composition. Such impacts of future changes in species communities should receive more attention in literature.     

Traffic noise induces oxidative stress and phytohormone imbalance in two urban plant species

Traffic noise pollution is one of the major environmental concerns in crowded cities worldwide. The objective of the present study was to investigate the effects of traffic noise on growth, hormonal balance, oxidative damage, and activity of antioxidant systems in two urban plant species, Tagetes patula and Salvia splendens. Each of the plant species were equally divided into 2 groups (control and traffic noise treatment) and each group was grown under identical controlled conditions in two separate growth chambers for two months. Traffic noise was recorded during peak traffic hours in a highly congested area of the city. Frequency analysis of traffic noise was conducted on the samples during the recording process. Plants in the traffic noise treatment group were exposed to 16 h of road traffic noise each day for a total of 15 days, while the control group was kept under complete silence. Traffic noise exposure led to significant decrease in growth indices of both plant species. The content of H₂O₂ and malondialdehyde (MDA) significantly increased upon traffic noise treatment in both species, followed by an increase in DPPH (2,2-diphenyl-1-picrylhydrazyl) anti-oxidant activity and the activity of antioxidant enzymes including catalase, peroxidase, and ascorbate peroxidase. Road traffic noise significantly reduced the content of phytohormones including zeatin, salicylic acid, indole-3-acetic acid, and gibberellic acid. On the other hand, the content of stress-related hormones including abscisic acid and jasmonic acid significantly increased in response to road traffic noise in both species. Based on our results, daily traffic noise can negatively affect growth and physiology of plants by inducing of oxidative damage and interfering with hormonal balance.     

Egg crypsis and clutch survival in three species of blackbirds (Icteridae)

For most open-nesting passerine birds, the primary function of egg pigmentation is crypsis. Interspecific variation for egg coloration is quite high, even among species that nest in the same locale. This variation might be maintained by selection to blend into species-specific nest cups, which vary in pattern because they are constructed from different materials. To test this hypothesis we photographed 30 clutches of Brewer's blackbirds ( Euphagus cyanocephalus ), 26 clutches of red-winged blackbirds ( Agelaius phoeniceus ), and 24 clutches of yellow-headed blackbirds ( Xanthocephalus xanthocephalus ), and quantified crypsis in terms of pattern (PD crypsis), brightness (MB crypsis) and brightness disparity (BD crypsis). The species were equally cryptic in terms of pattern crypsis, but differed significantly ( P < 0.01) in MB and BD crypsis. Intraspecific variation in pattern crypsis was surprisingly high. Mean clutch pattern disparity was highly correlated with mean background pattern disparity ( r = 0.999); this suggests selection for disparity matching, rather than for crypsis. We also monitored 25 clutches (7 Brewer's, 9 red-winged, and 9 yellow-headed blackbirds) for a 9-day period to determine if survival was related to crypsis. For Brewer's and yellow-headed blackbirds, successful nests were no more cryptic than failed nests. For red-winged blackbirds, there was a tendency ( P = 0.1) for less cryptic clutches to have higher survival; in addition, successful red-winged blackbird clutches had significantly ( P < 0.05) higher egg pattern disparity and background pattern disparity. Red-winged blackbird clutches are not concealed by overhanging vegetation, and high pattern disparity may enable them to blend into a large-scale heterogeneous background.     

Functional trait changes in the floras of 11 cities across the globe in response to urbanization

Urbanization causes major environmental changes globally, which can potentially homogenize biota across cities through the loss and gain of particular types of species. We examine whether urban environments consistently select for plants with particular traits and the implications of such changes on the functional composition of urban floras. We classified plant recorded in 11 cities around the globe as species that have either colonized (arrived and naturalized), persisted or been lost (local extirpation) following urbanization. We analyzed how 10 traits previously linked with plant responses to environmental conditions explained membership of these three groups, by comparing colonisers with persistent and extirpated plants through individual city‐level Bayesian models. Then, we used meta‐analysis to assess consistency of traits across urban areas. Finally, we explored several possible scenarios of functional change using these results. On average, urban colonizers had heavier seeds, unspecialised nutrient requirements, were taller and were annual species more often, especially when compared to locally extirpated plants. Common trends of functional change in urban plant communities include shifts towards taller and heavier‐seeded plants, and an increased prevalence of the short‐lived species, and plants without mutualistic nutritional strategies. Our results suggest that plant traits influence the species that succeed in urban environments worldwide. Different species use different ecological strategies to live in urban environments, as suggested by the importance of several traits that may appear as trait constellations. Plant height and seed mass were the only traits associated with both colonizer and extirpated plant status in urban environments. Based on our data, predicting colonization in urban environments may be easier than identifying extirpation‐prone plants; albeit some regional variation, colonization seems strongly driven by environmental conditions common to most cities (e.g. altered disturbance regimes), whereas extirpation may depend more on processes that vary across cities.     

Life history consequences of developing in anthropogenic noise

When environments change rapidly, adaptive phenotypic plasticity can ameliorate negative effects of environmental change on survival and reproduction. Recent evidence suggests, however, that plastic responses to human‐induced environmental change are often maladaptive or insufficient to overcome novel selection pressures. Anthropogenic noise is a ubiquitous and expanding disturbance with demonstrated effects on fitness‐related traits of animals like stress responses, foraging, vigilance, and pairing success. Elucidating the lifetime fitness effects of noise has been challenging because longer‐lived vertebrate systems are typically studied in this context. Here, we follow noise‐stressed invertebrates throughout their lives, assessing a comprehensive suite of life history traits, and ultimately, lifetime number of surviving offspring. We reared field crickets, Teleogryllus oceanicus, in masking traffic noise, traffic noise from which we removed frequencies that spectrally overlap with the crickets’ mate location song and peak hearing (nonmasking), or silence. We found that exposure to masking noise delayed maturity and reduced adult lifespan; crickets exposed to masking noise spent 23% more time in juvenile stages and 13% less time as reproductive adults than those exposed to no traffic noise. Chronic lifetime exposure to noise, however, did not affect lifetime reproductive output (number of eggs or surviving offspring), perhaps because mating provided females a substantial longevity benefit. Nevertheless, these results are concerning as they highlight multiple ways in which traffic noise may reduce invertebrate fitness. We encourage researchers to consider effects of anthropogenic disturbance on growth, survival, and reproductive traits simultaneously because changes in these traits may amplify or nullify one another.     

Acoustic and Temporal Partitioning of Cicada Assemblages in City and Mountain Environments

Comparing adaptations to noisy city environments with those to natural mountain environments on the community level can provide significant insights that allow an understanding of the impact of anthropogenic noise on invertebrates that employ loud calling songs for mate attraction, especially when each species has its distinct song, as in the case of cicadas. In this study, we investigated the partitioning strategy of cicada assemblages in city and mountain environments by comparing the acoustic features and calling activity patterns of each species, recorded using automated digital recording systems. Our comparison of activity patterns of seasonal and diel calling revealed that there was no significant temporal partitioning of cicada assemblages in either environment. In addition, there was no correlation between the acoustic distance based on spectral features and temporal segregation. Heterospecific spectral overlap was low in both city and mountain environments, although city and mountain cicada assemblages were subject to significantly different levels of anthropogenic or interspecific noise. Furthermore, for the common species found in both environments, the calling activity patterns at both seasonal and diel time scales were significantly consistent across sites and across environments. We suggest that the temporal calling activity is constrained by endogenous factors for each species and is less flexible in response to external factors, such as anthropogenic noise. As a result, cicada assemblages in city environments with low species diversity do not demonstrate a more significant temporal partitioning than those in mountain environments with high species diversity.     

How do invasive species travel to and through urban environments?

Globalisation has resulted in the movement of organisms outside their natural range, often with negative ecological and economic consequences. As cities are hubs of anthropogenic activities, with both highly transformed and disturbed environments, these areas are often the first point of entry for alien species. We compiled a global database of cities with more than one million inhabitants that data had on alien species occurrence. We then identified the most prominent pathways of introduction and vectors of spread of alien species in these cities. Most species were intentionally introduced to cities and were released or escaped from confinement. The majority of alien species then spread within cities through natural means (primarily unaided dispersal). Pathway prominence varied across the taxonomic groups of alien species: the most prominent pathway for plants and vertebrates was the escape pathway; for invertebrates the stowaway and contaminant pathways were most likely to facilitate introductions. For some organisms, pathway prominence varied with the geographical and climatic characteristics of the city. The characteristics of the cities also influenced the prominence of vectors of spread for alien species. Preventing the natural spread of alien species within cities, and into adjacent natural environments will be, at best, difficult. To prevent invasions, both the intentional and unintentional introduction of potentially harmful alien species to cities must be prevented. The pathways of introduction and vectors of spread identified here should be prioritised for management.     

Independent colonization of multiple urban centres by a formerly forest specialist bird species

Urban areas are expanding rapidly, but a few native species have successfully colonized them. The processes underlying such colonization events are poorly understood. Using the blackbird Turdus merula, a former forest specialist that is now one of the most common urban birds in its range, we provide the first assessment of two contrasting urban colonization models. First, that urbanization occurred independently. Second, that following initial urbanization, urban-adapted individuals colonized other urban areas in a leapfrog manner. Previous analyses of spatial patterns in the timing of blackbird urbanization, and experimental introductions of urban and rural blackbirds to uncolonized cities, suggest that the leapfrog model is likely to apply. We found that, across the western Palaearctic, urban blackbird populations contain less genetic diversity than rural ones, urban populations are more strongly differentiated from each other than from rural populations and assignment tests support a rural source population for most urban individuals. In combination, these results provide much stronger support for the independent urbanization model than the leapfrog one. If the former model predominates, colonization of multiple urban centres will be particularly difficult when urbanization requires genetic adaptations, having implications for urban species diversity.     

White‐crowned sparrow males show immediate flexibility in song amplitude but not in song minimum frequency in response to changes in noise levels in the field

The soundscape acts as a selective agent on organisms that use acoustic signals to communicate. A number of studies document variation in structure, amplitude, or timing of signal production in correspondence with environmental noise levels thus supporting the hypothesis that organisms are changing their signaling behaviors to avoid masking. The time scale at which organisms respond is of particular interest. Signal structure may evolve across generations through processes such as cultural or genetic transmission. Individuals may also change their behavior during development (ontogenetic change) or in real time (i.e., immediate flexibility). These are not mutually exclusive mechanisms, and all must be investigated to understand how organisms respond to selection pressures from the soundscape. Previous work on white‐crowned sparrows (Zonotrichia leucophrys) found that males holding territories in louder areas tend to sing higher frequency songs and that both noise levels and song frequency have increased over time (30 years) in urban areas. These previous findings suggest that songs are changing across generations; however, it is not known if this species also exhibits immediate flexibility. Here, we conducted an exploratory, observational study to ask whether males change the minimum frequency of their song in response to immediate changes in noise levels. We also ask whether males sing louder, as increased minimum frequency may be physiologically linked to producing sound at higher amplitudes, in response to immediate changes in environmental noise. We found that territorial males adjust song amplitude but not minimum frequency in response to changes in environmental noise levels. Our results suggest that males do not show immediate flexibility in song minimum frequency, although experimental manipulations are needed to test this hypothesis further. Our work highlights the need to investigate multiple mechanisms of adaptive response to soundscapes.     

Red-winged blackbird predation on periodical cicadas (Cicadidae: Magicicada spp.): bird behavior and cicada responses

Summary Predation by red-winged blackbirds Agelaius phoeniceus L. on 13-year periodical cicadas (Magicicada spp.) and reactions by periodical cicadas to predators were studied during emergence of Brood XIX during summer of 1985 in northwestern Arkansas (USA). Emergences of periodical cicadas are classic examples of predator satiation due to high local densities of cicadas and birds are the major predators of adult periodical cicadas. Reactions of periodical cicadas to predators were assessed by recording behaviors exhibited by cicadas when approached in trees by a human hand during the 3-week period of peak adult densitics. Most male cicadas made a noise in association with escape behaviors when approached, and 50% of the females, which are silent, attempted escape behaviors. Observations of predation attempts on cicadas by red-winged blackbirds were made during the period of peak predation pressure. Red-winged blackbirds spent less time searching for cicadas over that 2-week period of increasing predation pressure, and became increasingly efficient at capturing cicadas. Handling time of cicadas by red-winged blackbirds increased by about 20 seconds over that period, as blackbirds spent more time consuming female cicadas. The flysquawk response, used only by male cicadas, was effective in deterring red-winged blackbirds; only 5% of the attacks by blackbirds were successful when that behavior was exhibited. All cicadas that remained motionless and silent when approached by blackbirds were captured and consumed. Because females remained inactive when approached more often than did males, blackbirds may have consumed more female cicadas. Changes that appeared in reactions of preiodical cicadas to the model predator and to the attacks of blackbirds reflect both changes in the sex ratio of the cicada populations and changes in behaviors of cicadas associated with mating and egg laying. The loud noise made by male periodical cicadas at mating centers did not appear to deter predation by blackbirds. Changes in the behavior of blackbirds that appeared to be in association with greater predation on female periodical cicadas relates directly to aspects of foraging theory, particularly predictions concerning more selective foraging during periods of abundant food resources.