Community convergence in bird song

Species in similar habitats are often similar in morphology or behaviour, attributed to adaptation to similar environmental selection pressures, sometimes mediated by competitive interactions. For passerine songs, similarity of phenotype in identical habitats and character displacement have been documented, the former due to adaptation to the acoustics of the habitat, and the latter due to competition for acoustic space among species. If these phenomena are widespread, they should lead to community convergence of bird songs. Here, we test if passerine communities in similar habitats converge in song attributes or in acoustic differentiation among species. We compared the songs of European and North American Mediterranean climate passerine communities in open and closed habitats. Song frequency varied across different habitats but not continents. This was independent of both phylogeny and body size, indicating community convergence due to acoustic adaptation, rather than species sorting or similarity as a by-product of another type of ecological convergence. We found little evidence for regular spacing in song features among species, as would be expected if acoustic competition shapes within-community structure. However, for one of five song components, the open habitat communities showed a similar distribution of phenotypes on each continent. The proportion of interspecific variation in song explained by these effects was small. The fact that songs are complex signals that vary in many dimensions may explain why competition for acoustic space seems to be of small importance in structuring songs in these passerine communities.     

Inter- and Intraspecific Variation in the Acoustic Habitats of Three Marmot Species

Closely related species often have remarkably different vocalizations. Some of the variation in acoustic structure may result from species adapting their calls to maximize transmission through their acoustic environments. We document the relative magnitude of inter- and intraspecific variation in acoustic transmission properties of the habitats of three closely related marmot species to study the relative importance that the acoustic environment may have played in selecting for species-specific marmot alarm calls. We used spectrogram correlation to quantify the degree to which pure tones and alarm calls changed as they were broadcast through marmot home ranges to describe the acoustic habitats of golden (M. candata aurea), yellow-bellied (M. flaviventris), and alpine (M. marmota L.) marmots. Species lived in quantifiably different acoustic habitats. One analysis partitioned variation between species and between marmot social groups (nested within species). We found significant interspecific variation in the acoustic transmission fidelity of the three species' habitats and insignificant intraspecific variation between social groups. Further analysis of a larger sample of alarm calls broadcast through golden marmot social groups found significant intraspecific variation. Interspecific variation greater than intraspecific variation suggests that variable acoustic habitats may be responsible for at least some of the interspecific variation in alarm call structure. This is the first study to use spectrogram correlation to describe habitat acoustics. We discuss aspects of the method that may be useful for others seeking to quantify habitat acoustics.     

Local richness and distribution of the lizard fauna in natural habitat mosaics of the Brazilian Cerrado

We investigate local lizard richness and distribution in central Brazilian Cerrado, harbouring one of the least studied herpetofaunas in the Neotropical region. Our results are based on standardized samplings at 10 localities, involving 2917 captures of 57 lizard species in 10 families. Local richness values exceeded most presented in earlier studies and varied from 13 to 28 species, with modal values between 19 and 28 species. Most of the Cerrado lizard fauna is composed of habitat‐specialists with patchy distributions in the mosaic of grasslands, savannas and forests, resulting in habitat‐structured lizard assemblages. Faunal overlap between open and forested habitats is limited, and forested and open areas may act as mutual barriers to lizard distribution. Habitat use is influenced by niche conservatism in deep lineages, with iguanians and gekkotans showing higher use of forested habitats, whereas autarchoglossans are richer and more abundant in open habitats. Contrary to trends observed in Cerrado birds and large mammals, lizard richness is significantly higher in open, interfluvial habitats that dominate the Cerrado landscape. Between‐localities variation in lizard richness seems tied to geographical distance, landscape history and phylogenetic constraints, factors operating in other well‐studied lizard faunas in open environments. Higher richness in dominant, open interfluvial habitats may be recurrent in Squamata and other small‐bodied vertebrates, posing a threat to conservation as these habitats are most vulnerable to the fast, widespread and ongoing process of habitat destruction in central Brazil.     

Population genetic structure of male black grouse (Tetrao tetrix L.) in fragmented vs. continuous landscapes

We investigated the association of habitat fragmentation with genetic structure of male black grouse Tetrao tetrix. Using 14 microsatellites, we compared the genetic differentiation of males among nine localities in continuous lowland habitats in Finland to the genetic differentiation among 14 localities in fragmented habitats in the Alps (France, Switzerland and Italy). In both areas, we found significant genetic differentiation. However, the average differentiation, measured as theta, was more than three times higher in the Alps than in Finland. The greater differentiation found in the Alps is probably due to the presence of mountain ridges rising above natural habitats of the species, which form barriers to gene flow, and to a higher influence of genetic drift resulting from lower effective sizes in highly fragmented habitats. The detection of isolation by distance in the Alps suggests that gene flow among populations does occur. The genetic variability measured as gene diversity HE and allelic richness A was lower in the Alps than in Finland. This could result from the higher fragmentation and/or from the fact that populations in the Alps are isolated from the main species range and have a lower effective size than in Finland. This study suggests that habitat fragmentation can affect genetic structure of avian species with relatively high dispersal propensities.     

Local habitat complexity correlates with song complexity in a vocally elaborate honeyeater

Song complexity is an important behavioural trait in songbirds, subject to sexual selection. Elucidation of intraspecific variation in song complexity can provide insights into its evolution. In this study, we investigated song complexity variation in tūī (Prosthemadera novaeseelandiae), a vocally complex songbird endemic to New Zealand. At two separate nature reserves, we recorded male songs in two habitat types: forest remnants with high habitat complexity, and open habitats with lower habitat complexity. Analyses indicated strong evidence that song complexity was higher in forest habitats. Possible explanations for this divergence include: (i) competition between individuals results in higher quality, dominant males with more complex songs occupying forest habitats, and less competitive males occupying open habitat zones; (ii) forest habitats provide more abundant resources therefore higher tūī density, resulting in more complex songs; and (iii) a higher abundance of food in dense forest habitats may reduce nutritional stress during development resulting in full development of song nuclei. However, these hypotheses on the drivers of habitat effects on tūī song complexity remain to be tested.     

Song parameters of the fuscous honeyeater Lichenostomus fuscus correlate with habitat characteristics in fragmented landscapes

Both avian abundance and species richness decline in response to habitat loss and fragmentation. Studying variation in bird song structure across modified landscapes can provide insights into the effects of habitat alterations on coherence of social interactions within populations. Here, we tested whether fragmentation or change of habitat quality within box‐ironbark forest of central Victoria impacted cultural connectivity and song characteristics in fuscous honeyeater, a declining common Australian bird. First, we tested whether geographic distance and/or spatially‐explicit landscape connectivity models can explain patterns of song similarity across fragmented landscapes. We found no evidence that distance or habitat fragmentation impacts the nature and transmission of fuscous honeyeater song, and concluded that acoustic connectivity at the scale of our study is high. Second, we tested whether variation in habitat quality explains variation in song characteristics. In accordance with acoustic adaptation to habitat structure, birds sang longer songs in sites with more large trees and produced longer common song elements in sites with greater tree height. However, the acoustic adaptation hypothesis cannot explain the finding that in less‐disturbed landscapes with higher tree‐cover birds sang songs (and song elements) with higher maximum frequency and wider frequency bandwidth. We also found that birds sing longer and more variable songs of wider frequency bandwidth in less disturbed sites with a greater number of large mature trees, which may represent better feeding resources. Our study suggests that changes in song structure with habitat degradation could signal disturbed population processess, such as changes in the acoustic communication among resident birds.     

Habitat-dependent song divergence in the little greenbul: an analysis of environmental selection pressures on acoustic signals

Abstract.— Bird song is a sexual trait important in mate choice and known to be shaped by environmental selection. Here we investigate the ecological factors shaping song variation across a rainforest gradient in central Africa. We show that the little greenbul ( Andropadus virens ), previously shown to vary morphologically across the gradient in fitness-related characters, also varies with respect to song characteristics. Acoustic features, including minimum and maximum frequency, and delivery rate of song notes showed significant differences between habitats. In contrast, we found dialectal variation independent of habitat in population-typical songtype sequences. This pattern is consistent with ongoing gene flow across habitats and in line with the view that song variation in the order in which songtypes are produced is not dependent on habitat characteristics in the same way physical song characteristics are. Sound transmission characteristics of the two habitats did not vary significantly, but analyses of ambient noise spectra revealed dramatic and consistent habitat-dependent differences. Matching between low ambient noise levels for low frequencies in the rainforest and lower minimal frequencies in greenbul songs in this habitat suggests that part of the song divergence may be driven by habitat-dependent ambient noise patterns. These results suggest that habitat-dependent selection may act simultaneously on traits of ecological importance and those important in prezygotic isolation, leading to an association between morphological and acoustic divergence. Such an association may promote assortative mating and may be a mechanism driving reproductive divergence across ecological gradients.     

Song Post Height in Relation to Predator Diversity and Urbanization

Birds may sing from positions in the vegetation (song posts) to allow efficient transmission of sexual and territorial vocal displays while simultaneously minimizing the risk of predation because of avian and mammalian predators. Because urban areas are deficient in specialized avian predators, but have many cats while the opposite is the case for nearby rural areas, urban birds should display higher in the vegetation. In a comparison of the abundance of predators in three cities (Oslo, Brønderslev, Orsay), I show that avian predators are more common in rural areas, while mammalian predators are more common in urban areas. Singing birds sang from higher positions in the vegetation of urban than nearby rural areas. Differences in song post heights between urban and rural areas were consistent among cities, suggesting inherent specific difference in microhabitat choice. Bird species that have become urbanized recently had similar song post heights in urban and rural habitats, while species that have been urbanized for a long time sang from relatively higher song posts in urban areas. These findings suggest that urban and rural birds differ in habitat use when singing. These differences in song post choice between urban and rural habitats may have a number of consequences for vocal displays in the two different habitats.     

Temporal stability and change in a microgeographical pattern of song variation in the rufous-collared sparrow

We studied the pattern of song variation in the rufous-collared sparrow, Zonotrichia capensis, in the 1987 and 2000 breeding seasons in a 7-km² area covering open and closed habitats. We measured 14 quantitative song variables over a total of 390 individuals and found consistent differences between habitats in both years. In particular, songs of individuals from closed habitat sites had trills with longer trill intervals and lower frequencies than those of individuals from open habitats. This pattern of variation is interpreted as a song cline that correlates with the environmental gradient. Although this cline was stable in location and shape, it also showed several differences between years. The songs recorded during 2000 had trills with longer trill intervals and lower minimum frequencies than those recorded during 1987. Thus, this study is the first to directly document temporal changes in song variables in a Z. capensis population. This change would be expected if all the habitats studied were more closed in 2000 than in 1987, but we did not find obvious differences in habitat features between seasons. However, this modification of song structure is compatible with alternative explanations, such as subtle differences in habitat characteristics, changes in climatic variables, a delayed effect of a past modification of the environment and neutrality of the change.     

Convergent maternal care strategies in ungulates and macropods

Mammals show extensive interspecific variation in the form of maternal care. Among ungulates, there is a dichotomy between species in which offspring follow the mother ("following" strategy) versus species in which offspring remain concealed ("hiding" strategy). Here we reveal that the same dichotomy exists among macropods (kangaroos, wallabies and allies). We test three traditional adaptive explanations and one new life history hypothesis, and find very similar patterns among both ungulates and macropods. The three traditional explanations that we tested were that a "following" strategy is associated with (1) open habitat, (2) large mothers, and (3) gregariousness. Our new life-history hypothesis is that a "following strategy" is associated with delayed weaning, and thus with the "slow" end of the slow-fast mammalian life-history continuum, because offspring devote resources to locomotion rather than rapid growth. Our comparative test strongly supports the habitat structure hypothesis and provides some support for this new delayed weaning hypothesis for both ungulates and macropods. We propose that sedentary young in closed habitats benefit energetically by having milk brought to them. In open habitats, predation pressure will select against hiding. Followers will suffer slower growth to independence. Taken together, therefore, our results provide the first quantitative evidence that macropods and ungulates are convergent with respect to interspecific variation in maternal care strategy. In both clades, differences between species in the form of parental care are due to a similar interaction between habitat, social behavior, and life history.     

Habitat selection for parasite-free space by hosts of parasitic cowbirds

Choice of breeding habitat can have a major impact on fitness. Sensitivity of habitat choice to environmental cues predicting reproductive success, such as density of harmful enemy species, should be favored by natural selection. Yet, experimental tests of this idea are in short supply. Brown-headed cowbirds Molothrus ater commonly reduce reproductive success of a wide diversity of birds by parasitizing their nests. We used song playbacks to simulate high cowbird density and tested whether cowbird hosts avoid such areas in habitat selection. Host species that made settlement decisions during manipulations were significantly less abundant in the cowbird treatment as a group. In contrast, hosts that settled before manipulations started and non-host species did not respond to treatments. These results suggest that hosts of cowbirds can use vocal cues to assess parasitism risk among potential habitat patches and avoid high risk habitats. This can affect community structure by affecting habitat choices of species with differential vulnerability.     

Species-specific song convergence in a moving hybrid zone between two passerines

Moving hybrid zones are receiving increasing attention. However, so far little is known about the proximate mechanisms underlying these movements. Signalling behaviour, by individuals engaged in interspecific sexual and aggressive interactions, may play a crucial role. In this study, we investigated song variation within a moving hybrid zone between two warblers, Hippolais polyglotta and H. icterina . In these species, song is involved in interspecific territoriality and, probably, in mixed pairings. We showed that allopatric populations of the two species are clearly acoustically differentiated. However, interspecific differences faded out in sympatry as a result of an overall pattern of convergence. Unexpectedly, the two species converged for different song parameters, namely temporal parameters for H. icterina and syntax for H. polyglotta . Hybridization and interspecific competition could explain convergence in H. icterina . Instead, in H. polyglotta we suggest that local adaptation to habitat and interspecific learning might contribute to convergence. We particularly stress that cross-species learning, by maintaining high levels of interspecific interactions, may influence the movement of the zone.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 507–517.     

Distribution and abundance of four caiman species (Crocodylia: Alligatoridae) in Jaú National Park,Amazonas, Brazil

Jaú National Park is a large rain forest reserve that contains small populations of four caiman species. We sampled crocodilian populations during 30 surveys over a period of four years in five study areas. We found the mean abundance of caiman species to be very low (1.0 +/- 0.5 caiman/km of shoreline), independent of habitat type (river, stream or lake) and season. While abundance was almost equal, the species' composition varied in different waterbody and study areas. We analysed the structure similarity of this assemblage. Lake and river habitats were the most similar habitats, and inhabited by at least two species, mainly Caiman crocodilus and Melanosuchus niger. However, those species can also inhabit streams. Streams were the most dissimilar habitats studied and also had two other species: Paleosuchus trigonatus and P. palpebrosus. The structure of these assemblage does not suggest a pattern of species associated and separated by habitat. Trends in species relationships had a negative correlation with species of similar size, C. crocodilus and P. trigonatus, and an apparent complete exclusion of M. niger and P. trigonatus. Microhabitat analysis suggests a slender habitat partitioning. P. trigonatus was absent from river and lake Igapó (flooded forest), but frequent in stream Igapó. This species was the most terrestrial and found in microhabitats similar to C. crocodilus (shallow waters, slow current). Melanosuchus niger inhabits deep, fast moving waters in different study areas. Despite inhabiting the same waterbodies in many surveys, M. niger and C. crocodilus did not share the same microhabitats. Paleosuchus palpebrosus was observed only in running waters and never in stagnant lake habitats. Cluster analysis revealed three survey groups: two constitute a mosaic in floodplains, (a) a cluster with both M. niger and C. crocodilus, and another (b) with only C. crocodilus. A third cluster (c) included more species, and the presence of Paleosuchus species. There was no significant difference among wariness of caimans between disturbed and undisturbed localities. However, there was a clear trend to increase wariness during the course of consecutive surveys at four localities, suggesting that we, more than local inhabitants, had disturbed caimans. The factors that are limiting caiman populations can be independent of human exploitation. Currently in Amazonia, increased the pressure of hunting, habitat loss and habitat alteration, and there is no evidence of widespread recovery of caiman populations. In large reserves as Jaú without many disturbance, most caiman populations can be low density, suggesting that in blackwater environments their recovery from exploitation should be very slow.     

No Habitat Selection during Spring Migration at a Meso-Scale Range across Mosaic Landscapes: A Case Study with the Woodcock (Scolopax rusticola)

Success of migration in birds in part depends on habitat selection. Overall, it is still poorly known whether there is habitat selection amongst landbird migrants moving across landscapes. Europe is chiefly covered by agro-forestry mosaic landscapes, so migratory species associated to either agricultural landscapes or woodland habitats should theoretically find suitable stopover sites along migration. During migration from wintering to breeding quarters, woodcocks (Scolopax rusticola) tagged with PTT satellite-tracking transmitters were used to test for the hypothesis that migrants associated to agro-forest habitats have no habitat selection during migration, at a meso-scale level. Using a GIS platform we extracted at a meso-scale range habitat cover at stopover localities. Results obtained from comparisons of soil covers between points randomly selected and true stopover localities sites revealed, as expected, the species may not select for particular habitats at a meso-scale range, because the habitat (or habitats) required by the species can be found virtually everywhere on their migration route. However, those birds stopping over in places richer in cropland or mosaic habitats including both cropland and forest and with proportionally less closed forest stayed for longer than in areas with lower surfaces of cropland and mosaic and more closed forest. This suggests that areas rich in cropland or mosaic habitat were optimal.     

Local extinctions and recolonisations of passerine bird populations in small woods

This paper considers, for eight species of woodland bird, the factors that influenced both local extinctions and recolonisations in 145 woods over 3 years. In all species, probability of local extinction was inversely related to population size; most local extinctions occurred in woods containing one to three breeding pairs. However, considerable variation in extinction probabilities occurred between species and between years. In addition, the suitability of habitat within a wood (more extinctions in less suitable woods) was important for wren Troglodytes troglodytes, song thrush Turdus philomelos and blue tit Parus caeruleus; also, the structure of the surrounding landscape was important for blue tit, great tit Parus major, and chaffinch Fringilla coelebs (more extinctions in localities with less woodland). In only two species was the probability of recolonisation related to any of the measured variables. Wrens were more likely to recolonise larger woods, whereas song thrushes were more likely to recolonise woods with a high habitat suitability rating and those which are more isolated from other woodland     

Habitat density, song structure and dialects in the Madagascar paradise flycatcher Terpsiphone mutata

Bird song is an important signal for long-distance intraspecific communication. Thus, song characteristics are expected to have undergone adaptive modification to maximise propagation through the environment. Acoustic properties of song should be correlated with the habitat in which they are broadcast, and neighbouring individuals should sing similar songs because they share the same local acoustic environment. We tested these predictions by investigating associations between habitat characteristics and song attributes in the Madagascar paradise flycatcher Terpsiphone mutata . We also examined whether song dialects were distributed non-randomly through the study site and which features of habitat density best explained these associations. We found that individuals on territories in denser habitat sang longer songs at lower frequencies. Features of song structure were related to both understorey and canopy cover density. The distribution of dialects within the study site was also strongly associated with variation in understorey cover. These findings suggest that habitat variation contributes to maintaining microgeographic variation in song structure in this species.     

Geographic variation in advertisement calls in a tree frog species: gene flow and selection hypotheses

Background

In a species with a large distribution relative to its dispersal capacity, geographic variation in traits may be explained by gene flow, selection, or the combined effects of both. Studies of genetic diversity using neutral molecular markers show that patterns of isolation by distance (IBD) or barrier effect may be evident for geographic variation at the molecular level in amphibian species. However, selective factors such as habitat, predator, or interspecific interactions may be critical for geographic variation in sexual traits. We studied geographic variation in advertisement calls in the tree frog Hyla japonica to understand patterns of variation in these traits across Korea and provide clues about the underlying forces for variation.

Methodology

We recorded calls of H. japonica in three breeding seasons from 17 localities including localities in remote Jeju Island. Call characters analyzed were note repetition rate (NRR), note duration (ND), and dominant frequency (DF), along with snout-to-vent length.

Results

The findings of a barrier effect on DF and a longitudinal variation in NRR seemed to suggest that an open sea between the mainland and Jeju Island and mountain ranges dominated by the north-south Taebaek Mountains were related to geographic variation in call characters. Furthermore, there was a pattern of IBD in mitochondrial DNA sequences. However, no comparable pattern of IBD was found between geographic distance and call characters. We also failed to detect any effects of habitat or interspecific interaction on call characters.

Conclusions

Geographic variations in call characters as well as mitochondrial DNA sequences were largely stratified by geographic factors such as distance and barriers in Korean populations of H. japoinca. Although we did not detect effects of habitat or interspecific interaction, some other selective factors such as sexual selection might still be operating on call characters in conjunction with restricted gene flow.     

Habitat Quality, Social Dominance and Dawn Chorus Song Output in Black-Capped Chickadees

Individuals that settle in poor habitats may reduce resource investments in various life history traits; for example, resources may be withheld from costly advertisement signals. There may be geographic variation in advertisement levels that correspond with habitat quality; however, this is poorly documented and it is unclear whether such habitat effects have consequences for the function of mate‐choice signals within habitats. We examined song output of male black‐capped chickadees (Poecile atricapillus) during the dawn chorus in two contrasting habitats (mature forest vs. young forest) known to differ in quality. Dominance rank is indicative of phenotypic quality in this species and was assessed during the preceding winter. We measured the song output of males participating in the dawn chorus during the nesting/egg laying period. Males living in young forest had reduced song output; however, a habitat–rank interaction term showed that dominant birds accounted for most of this difference. This suggests that signal reliability might be lost in poor habitats. We generate several hypotheses to explain these findings.     

Multi‐scale effects of habitat structure and landscape context on a vertebrate with limited dispersal ability (the brown‐throated sloth,Bradypus variegatus)

As human population, food consumption, and demand for forest products continue to rise over the next century, the pressures of land‐use change on biodiversity are projected to intensify. In tropical regions, countryside habitats that retain abundant tree cover and structurally complex canopies may complement protected areas by providing suitable habitats and landscape connectivity for a significant portion of the native biota. Species with low dispersal capabilities are among the most at risk of extinction as a consequence of land‐use change. We assessed how the spatial distribution of the brown‐throated sloth (Bradypus variegatus), a model species for a vertebrate with limited dispersal ability, is shaped by differences in habitat structure and landscape patterns of countryside habitats in north‐central Costa Rica using a multi‐scale framework. We quantified the influence of local habitat characteristics and landscape context on sloth occurrence using mixed‐effects logistic regression models. We recorded 27 sloths within countryside habitats and found that both local and landscape factors significantly influenced their spatial distribution. Locally, sloths favored structurally complex habitats, with greater canopy cover and variation in tree height and basal area. At the landscape scale, sloths demonstrated a preference for habitats with high proportions of forest and nearby large tracts of forest. Although mixed‐use areas and tree plantations are not substitutes for protected forests, our results suggest they provide important supplemental habitats for sloths. To promote the conservation and long‐term viability of sloth populations in the tropical countryside, we recommend that land managers retain structurally complex vegetation and large patches of native habitat.     

Interspecific variation in plumage colour among birds: species recognition or light environment?

Abstract The traditional explanation for interspecific plumage colour variation in birds is that colour differences between species are adaptations to minimize the risk of hybridization. Under this explanation, colour differences between closely related species of birds represent reproductive character displacement. An alternative explanation is that interspecific variation in plumage colour is an adaptive response to variation in light environments across habitats. Under this explanation, differences in colour between closely related species are a product of selection on signal efficiency. We use a comparative approach to examine these two hypotheses, testing the effects of sympatry and habitat use, respectively, on divergence in male plumage colour. Contrary to the prediction of the Species Isolation Hypothesis, we find no evidence that sympatric pairs of species are consistently more divergent in coloration than are allopatric pairs of species. However, in agreement with the Light Environment Hypothesis, we find significant associations between plumage coloration and habitat use. All of these results remain qualitatively unchanged irrespective of the statistical methodology used to compare reflectance spectra, the body regions used in the analyses, or the exclusion of areas of plumage not used in sexual displays. Our results suggest that, in general, interspecific variation in plumage colour among birds is more strongly influenced by the signalling environment than by the risk of hybridization.