Diverge or merge? The effect of sympatric occurrence on the territorial vocalizations of the vinaceous dove Streptopelia vinacea and the ring‐necked dove S. capicola

Sympatric occurrence of two closely related species is expected to lead to diverging or converging shifts in signal characteristics of one or both species. We examined signal characteristics in the vinaceous dove Streptopelia vinacea and the ring-necked dove S. capicola , two sister species that are mainly allopatric but occur in sympatry in northwestern Uganda. Vocal characteristics of the birds in sympatry were compared with those of an adjacent and a distant allopatric population of each species. The sympatric population showed intermediate values between the allopatric populations from Uganda. However, within each species there is little geographic variation between distant allopatric populations. Since vocal differences between dove species have a genetic base, the convergence in vocal characteristics is most likely explained by hybridization. Probably, the two species came into secondary contact relatively recently. Climatic changes during the last several thousand years and recent habitat changes caused by the growing human population, may have allowed Streptopelia capicola to extend its range in the northern direction.     

Male territorial vocalizations and responses are decoupled in an avian hybrid zone

A core area of speciation research concerns the coevolution of species-specific signals and the selective sensitivity to such signals. Signals and responses to them should be tuned to each other, to be effective in intraspecific communication. Hybrid zones are ideal to study the presence of such 'behavioural coupling' and the mechanisms governing it, and this has rarely been done. Our study examines acoustic signals of males and their response to them in the context of territorial interactions in a natural hybrid zone between two dove species, Streptopelia vinacea and Streptopelia capicola. Male signals are important in hybrid zone dynamics as they are essential for territory establishment, which is crucial for successful reproduction. We tested whether the response of individual male hybrids is linked to how similar their own signal is to the playback signal. We did not find evidence for behavioural coupling. The combined evidence from the low level of response to hybrid and heterospecific signals outside the hybrid zone and a lack of coupling within the hybrid zone suggests that perceptual learning may explain our results. Learning to respond to locally abundant signals may be the best individual strategy and is likely to contribute to the maintenance of a hybrid zone.     

Hybrid vocalizations are effective within, but not outside, an avian hybrid zone

Secondary contact between closely related species can lead tohybridization. The fitness of hybrid individuals within andoutside the hybrid zone determines whether the hybrid zone expandsinto the ranges of the 2 parental species or remains a stable,geographically narrow area in between the allopatric rangesof the parental species. In birds, vocalizations play an importantrole in male–male competition and female mate choice andare often affected by hybridization. One of the factors thatwill influence male hybrid fitness is the ability to defenda territory against competitors by vocalizing. We tested theefficacy of territorial signals of hybrids of 2 dove species,Streptopelia vinacea and Streptopelia capicola, compared withthe vocalizations of the parental species. With playback experiments,we assessed the response to hybrid and the 2 parental speciesvocalizations in the hybrid zone and adjacent allopatric populationsof each species. In the hybrid zone, males did not respond differentlyto the 3 vocalization types. In both allopatric populations,however, males responded more to conspecific than to heterospecificsignals and the response strength to hybrid signals was intermediate.Therefore, in the allopatric populations, hybrid males may havea reduced success in defending territories. In male–maleinteractions in the hybrid zone, hybrids may not have a disadvantagecompared with males of the parental species. The ability todefend a territory against competitors may thus help maintaina stable hybrid zone in the area of overlap.     

Molecular classification of Pakistani collared dove through DNA barcoding

Pakistan is bestowed by a diversified array of wild bird species including collared doves of which the taxonomy has been least studied and reported. DNA barcoding is a geno-taxonomic tool that has been used for characterization of bird species using mitochondrial cytochrome c oxidase I gene (COI). This study aimed to identify taxonomic order of Pakistani collared dove using DNA barcoding. Purposely herein, we present a phylogenetic analysis of Pakistani collared dove based on 650 base pairs of COI gene sequences. Analysis of phylogenetic tree revealed that Pakistani collared dove shared a common clade with Eurasian collared dove (Streptopelia decaocto) and African collared dove (Streptopelia roseogrisea) which indicated a super-species group in Streptopelia genus. This is the first report of molecular classification of Pakistani collared dove using DNA barcoding.     

Collared Dove Responses to Playback: Slaves to the Rhythm

Temporal features are the most important parameters for differentiating among the perch coo vocalizations of doves in the genus Streptopelia . The role of temporal features in inducing territorial responses was studied for one of these species, the collared dove ( S. decaocto ), using playback experiments. The playback stimuli consisted of manipulated coos, which were tested against a preplayback period and against unmanipulated control coos. In a first set of three manipulations, the temporal structure was altered by removing an element from the species-typical three-element coos. A coo lacking the third element, which is a naturally occurring variant, was as equally effective as the control in eliciting a territorial response. The same applied to a coo lacking the second and longest element. In contrast, a coo lacking the first element did not elicit a significant increase in territorial activities compared with the preplayback period, and there was a significant reduction in response compared with the control coo. A second set of two manipulations concerned changes in the species-typical rhythm. The two variants both led to a significant reduction in response compared with the normal rhythm, although they were still effective in eliciting an increase in activities compared with the preplayback period. The results indicate that the rhythm of territorial cooing plays an important role in communicating to conspecifics. As different sympatrically living dove species differ in particular in temporal features, rhythm may be an important cue for species recognition and may contribute to reproductive isolation.     

Reproductive character displacement and signal ontogeny in a sympatric assemblage of electric fish

The reproductive signals of two or more taxa may diverge in areas of sympatry, due to selection against costly reproductive interference. This divergence, termed reproductive character displacement (RCD), is expected in species-rich assemblages, where interspecific signal partitioning among closely related species is common. However, RCD is usually documented from simple two-taxon cases, via geographical tests for greater divergence of reproductive traits in sympatry than in allopatry. We propose a novel approach to recognizing and understanding RCD in multi-species communities--one that traces the displacement of signals within multivariate signal space during the ontogeny of individual animals. We argue that a case for RCD can be made if the amount of signal displacement between a pair of species after maturation is negatively correlated to distance in signal space before maturation. Our application of this approach, using a dataset of communication signals from a sympatric Amazonian assemblage of the electric fish genus Gymnotus, provides strong evidence for RCD among multiple species. We argue that RCD arose from the costs of heterospecific mismating, but interacted with sexual selection--favoring the evolution of conspicuous male signals that not only serve for mate-choice, but which simultaneously facilitate species recognition.     

Degree of male ornamentation affects female preference for conspecific versus heterospecific males.

Several studies have shown female preference for conspecific males with the attached artificial ornaments of more elaborate heterospecifics. However, preference for heterospecifics under natural conditions is relatively rare. We tested what factors affect behavioural mechanisms of species isolation using three species of estrildid finch (genus Uraeginthus) that occur in both sympatry and allopatry. These finches differ in degree of sexual dimorphism; male ornamentation; behavioural and morphological similarity; and phylogenetic distance. Paired mate-choice trials were used in which females were presented with a conspecific and heterospecific male to test which of the above between-species differences best predicted the degree of premating isolation. The three species differed in the degree of species-specific mate preference shown. Females from the brighter two species discriminated against dull males, independently of sympatry-allopatry, similarity and phylogenetic distance. Females from the dull species reacted to conspecific males and brighter heterospecific males equally strongly, independently of similarity and phylogenetic distance. In contrast to previous studies, an equal preference for heterospecific and conspecific males was found under natural conditions. It is suggested that differences between closely related species in male ornamentation affect the likelihood that premating isolation will occur due to the fact that sexual selection tends to drive preferences for exaggerated ornamentation.     

Alarm call‐based discrimination between common cuckoo and Eurasian sparrowhawk in a Chinese population of great tits

Morphological resemblance of the common cuckoo Cuculus canorus to the Eurasian sparrowhawk Accipiter nisus has been regarded as an example of predator mimicry. Common hosts could distinguish parasites as the result of coevolution, while rare hosts or non‐hosts may mistake cuckoos for hawks because rare hosts or non‐hosts behave similarly when faced with these two species. Birds usually produce alarm calls in addition to showing behavioral responses when in danger. However, previous studies of identification by rare hosts or non‐hosts of sparrowhawks usually lacked experimental evidence of alarm calls. Great tits Parus major, a rare cuckoo host, perform similar behaviors and usually produce alarm calls in response to sparrowhawks and common cuckoos. Here, we tested whether great tits could distinguish common cuckoo from sparrowhawk based on analysis of their alarm calls and the effects of playback of alarm calls on conspecific behavior. Previous studies showed that great tits have a complex communication system that conveys information about predators, and they could perform different kinds of response behavior to different alarm calls. If great tits have not made the ability to distinguish between common cuckoo and sparrowhawk, then their acoustic responses to these two species and their response behaviors in playback experiments should be similar. Specimens of a common cuckoo (parasite), a sparrowhawk (predator) and an Oriental turtle dove Streptopelia orientalis (harmless control) were used to elicit and subsequently record the response behavior and alarm calls of great tits. There was no significant difference in behavioral response among great tits when exposed to the dummy of cuckoo, sparrowhawk and dove. In contrast, they differed significantly in alarm calls. Great tits produced more notes per call that contained increasing D‐type and decreasing I‐type notes when responding to sparrowhawk as compared to cuckoo or dove. In playback experiments, we found that great tits responded more strongly to great tit hawk than to great tit cuckoo or great tit dove alarm calls. Our study suggests that great tits are able to distinguish sparrowhawks from common cuckoos and convey relevant information in alarm calls by adjusting the number and combinations of notes of a single call type.     

Divergence in mating signals correlates with genetic distance and behavioural responses to playback

Animals use acoustic signals to defend resources against rivals and attract breeding partners. As with many biological traits, acoustic signals may reflect ancestry; closely related species often produce more similar signals than do distantly related species. Whether this similarity in acoustic signals is biologically relevant to animals is poorly understood. We conducted a playback experiment to measure the physical and vocal responses of male songbirds to the songs of both conspecific and allopatric‐congeneric animals that varied in their acoustic and genetic similarity. Our subjects were territorial males of four species of neotropical Troglodytes wrens: Brown‐throated Wrens (Troglodytes brunneicollis), Cozumel Wrens (T. beani), Clarion Wrens (T. tanneri) and Socorro Wrens (T. sissonii). Our results indicate that birds respond to playback of both conspecific and allopatric‐congeneric animals; that acoustic differences increase with genetic distance; and that genetic divergence predicts the strength of behavioural responses to playback, after removing the effects of acoustic similarity between subjects’ songs and playback stimuli. Collectively, these results demonstrate that the most distantly related species have the most divergent songs; that male wrens perceive divergence in fine structural characteristics of songs; and that perceptual differences between species reflect evolutionary history. This study offers novel insight into the importance of acoustic divergence of learned signals and receiver responses in species recognition.     

Sympatry drives colour and song evolution in wood-warblers (Parulidae)

Closely related species often exhibit similarities in appearance and behaviour, yet when related species exist in sympatry, signals may diverge to enhance species recognition. Prior comparative studies provided mixed support for this hypothesis, but the relationship between sympatry and signal divergence is likely nonlinear. Constraints on signal diversity may limit signal divergence, especially when large numbers of species are sympatric. We tested the effect of sympatric overlap on plumage colour and song divergence in wood-warblers (Parulidae), a speciose group with diverse visual and vocal signals. We also tested how number of sympatric species influences signal divergence. Allopatric species pairs had overall greater plumage and song divergence compared to sympatric species pairs. However, among sympatric species pairs, plumage divergence positively related to the degree of sympatric overlap in males and females, while male song bandwidth and syllable rate divergence negatively related to sympatric overlap. In addition, as the number of species in sympatry increased, average signal divergence among sympatric species decreased, which is likely due to constraints on warbler perceptual space and signal diversity. Our findings reveal that sympatry influences signal evolution in warblers, though not always as predicted, and that number of sympatric species can limit sympatry''s influence on signal evolution.     

Ecology,song similarity and phylogeny predict natural hybridization in an avian family

Hybridization is important in the evolution of many animal groups; however, broad scale patterns of natural hybridization are still poorly understood. Using phylogenetic comparative analyses, we tested for relationships between demographic, ecological, phenotypic and phylogenetic variables and the incidence of natural hybridization among 45 species of North American wood warbler. Since 1980, hybrids have been documented in 24 species (53 %). We detected negative relationships between the incidence of hybridization and both breeding range size and phylogenetic distance, and positive relationships between the incidence of hybridization and (1) song similarity, (2) the extent of breeding sympatry of species pairs and (3) the number of additional heterospecific warbler species that co-occur during breeding. Neither population size nor breeding habitat quality (as measured by threats to survival or reproduction in breeding areas) explained variation in the incidence of hybridization. Our results suggest a potential role of limited breeding habitat in wood warbler hybridization events and a net positive effect of breeding sympatry. They also support the hypothesis that mating signal similarity facilitates hybridization events and are consistent with an increase in reproductive isolation with increasing genetic divergence. Our results also suggest the presence of phylogenetic signal in wood warbler hybridization. By investigating natural hybridization at the taxonomic level of family, we have identified several previously undocumented patterns of natural hybridization. This study demonstrates the utility of examining the combined effects of ecology, demography, phenotype and phylogeny when studying variation in the expression of natural hybridization among taxa.     

Sexual differentiation and seasonal variation in response to conspecific and heterospecific acoustic signals

Interspecific territoriality is frequently reported between closely related species; however, few studies have demonstrated interspecific territoriality between distantly related species living in sympatry. We conducted playback experiments to investigate territorial behaviour in male and female White‐bellied Wrens (Uropsila leucogastra) in response to simulated conspecific and heterospecific intruders during the breeding and non‐breeding seasons. We explored whether heterospecific songs of the Happy Wren (Pheugopedius felix), a distantly related species and ecological competitor, elicited antagonistic responses from focal White‐bellied Wrens, and whether such responses differed between the sexes. We also examined whether male and female responses to conspecific and heterospecific rivals varied with season. We found that male White‐bellied Wrens always responded to conspecific song, and responded significantly more to heterospecific song compared to a control stimulus (Tropical Parula, Setophaga pitiayumi). In contrast, although female White‐bellied Wrens responded strongly to conspecific song, their response to heterospecific song did not differ significantly from the control stimulus. The proportion of males that responded to heterospecific songs and the proportion of females that responded to conspecific songs varied seasonally, showing significantly lower responses during the breeding season. The intense responses of male White‐bellied Wrens to playback of heterospecific songs suggest that they recognise ecological competitors based on their vocal signals. Furthermore, the decrease in agonistic interactions during the breeding season is in line with the hypothesis that aggressive behaviour may be detrimental to reproductive and parental activity, and the hypothesis that heterospecific animals pose less of a threat during the breeding season.     

Co-Occurrence and Occupancy of Mourning Doves and Eurasian Collared-Doves

Understanding how land cover and potential competition with invasive species shape patterns of occupancy, extirpation, and colonization of native species across a landscape can help target management for declining native populations. Mourning dove (Zenaida macroura) populations have declined throughout the United States from 1965–2015. The expansion of the Eurasian collared-dove (Streptopelia decaocto), an introduced species with similar food preferences, may further threaten mourning dove populations. We analyzed data from 2009–2016 from a large-scale monitoring program in the Western Great Plains of the United States in a 2-species occupancy model to assess the effects of collared-doves on mourning dove distributions, while accounting for imperfect detection and variation in land cover across the landscape. Mourning dove occupancy was stable or increasing across our study area, and despite overlap in resource use and co-occurrence between mourning doves and Eurasian collared-doves, we found no evidence that collared-doves are extirpating mourning doves from preferred habitat during the breeding season. © 2020 The Wildlife Society.     

Manifold influences of phylogenetic structure on a plant–herbivore network

Ecologists are increasingly aware of the interplay between evolutionary history and ecological processes in shaping current species interaction patterns. The inclusion of phylogenetic relationships in studies of species interaction networks has shown that closely related species commonly interact with sets of similar species. Notably, the degree of phylogenetic conservatism in antagonistic ecological interactions is frequently stronger among species at lower trophic levels than among those at higher trophic levels. One hypothesis that accounts for this asymmetry is that competition among consumer species promotes resource partitioning and offsets the maintenance of dietary similarity by phylogenetic inertia. Here, we used a regional plant–herbivore network comprised of Asteraceae species and flower‐head endophagous insects to evaluate how the strength of phylogenetic conservatism in species interactions differs between the two trophic levels. We also addressed whether the asymmetry in the strength of the phylogenetic signal between plants and animals depends on the overall degree of relatedness among the herbivores. We show that, beyond the previously reported compositional similarity, closely related species also share a greater proportion of counterpart phylogenetic history, both for resource and consumer species. Comparison of the patterns found in the entire network with those found in subnetworks composed of more phylogenetically restricted groups of herbivores provides evidence that resource partitioning occurs mostly at deeper phylogenetic levels, so that a positive phylogenetic signal in antagonist similarity is detectable even between closely related consumers in monophyletic subnetworks. The asymmetry in signal strength between trophic levels is most apparent in the way network modules reflect resource phylogeny, both for the entire network and for subnetworks. Taken together, these results suggest that evolutionary processes, such as phylogenetic conservatism and independent colonization history of the insect groups may be the main forces generating the phylogenetic structure observed in this particular plant–herbivore network system.     

Invasive birds in a novel landscape: habitat associations and effects on established species

Studying the relationships among introduced species, their preferred habitats, and native species can be important for predicting the effects of invasions on native populations. Examining the colonization of North America by the Eurasian collared‐dove Streptopelia decaocto, we quantified the habitat characteristics of sites most likely to be occupied by this invasive bird species in the early stages of the invasion. Further, we studied the relationship between collared‐dove abundance and the abundance of other dove species in the study area, anticipating a negative effect on established species following the introduction of a potential competitor. Linking satellite‐derived landcover data with winter bird community data gathered from 444 study sites in Florida, USA from 1999 to 2008, we found that collared‐doves were more likely to occur in landscapes that had been highly‐modified by human activity than in forested landscapes. Collared‐dove abundance increased as the proportion of the landscape characterized as low‐intensity development and medium/high‐intensity development increased. The probability of collared‐doves occurring at a site was also related to the spatial proximity of other sites reporting doves (positive spatial autocorrelation). Contrary to our expectations, the site‐level abundance of four other dove species all increased with collared‐dove abundance throughout the sampling period. Interactions between collared‐doves and native species should be further studied in different environments as this invasive bird rapidly colonizes North America.     

Social aspects of the termination of incubation behaviour in the ring dove (Streptopelia risoria)

The role of male and female ring doves (Streptopelia risoria) in the termination of incubation behaviour was studied first by observing heterosexual pairs (male and female) and homosexual pairs (female and female) (experiment 1). Male/female pairs incubate for 22 days whether squab or unhatched eggs are in the nest; female/female pairs incubate for 25 days or more. Males take the lead in terminating incubation. Another group of doves, separated from their (heterosexual) partners late in incubation, were exposed to one of four stimulus conditions: (a) an incubating dove; (b) a reproductively-inactive dove; (c) a reproductively-active dove; (d) visual isolation. Females extended the duration of their incubation when exposed to the sight of another dove, except when the stimulus dove was reproductively active. Males did not extend their incubation in response to these social cues. These social stimuli terminating incubation behaviour coordinate the withdrawal from the nest of male and female in preparation for the next breeding episode in this monogamous species.     

Collared doves Streptopelia decaocto display with high, near-maximal muscle powers, but at low energetic cost

Display flight in collared doves Streptopelia decaocto consists of a rapid ascent from a perch, followed by a glide back to the same or near-by perch. The hypothesis that ascending flight presents an honest signal of the maximum power generating capacity of the displayer is considered. Using a deliberately conservative approach to calculating the power requirements of collared dove display flights, the highest power performance in an ascending collared dove is equivalent to that derived from wind tunnel studies on the morphologically similar ringed turtle-dove Streptopelia risoria . Muscle power requirements are very high (up to 232  W/kg), representing the highest performance measured in the field to date. However, the maximum energy requirements over an hour due to climbing power equates to only 5% of the basal metabolic rate. Display flight in collared doves appears to be an efficient and 'uncheatable' system for demonstrating fitness, either for mating or for territory defence.     

Closely related species of birds differ more in body size when their ranges overlap—in warm,but not cool,climates

Differences in body size are widely thought to allow closely related species to coexist in sympatry, but body size also varies as an adaptive response to climate. Here, we use a sister lineage approach to test the prediction that body size differences between closely related species of birds worldwide are greater for species whose ranges are sympatric rather than allopatric. We further test if body size differences among sympatric versus allopatric species vary with geography, evolutionary distance, and environmental temperatures. We find greater differences in size among sympatric compared with allopatric lineages, but only in closely related species that live where mean annual temperatures are above 25°C. These size differences in warm environments declined with the evolutionary distance between sister lineages. In species living in cooler regions, closely related allopatric and sympatric species did not differ significantly in size, suggesting either that colder temperatures constrain the evolutionary divergence of size in sympatry, or that the biotic selective pressures favoring size differences in sympatry are weaker in colder environments. Our results are consistent with suggestions by Wallace, Darwin, and Dobzhansky that climatic selective pressures are more important in cooler environments (e.g., high elevations and latitudes) whereas biotic selective pressures dominate in warm environments (e.g., lowland tropics).     

Reproductive character displacement of female,but not male song discrimination in an avian hybrid zone

Divergence of male sexual signals and female preferences for those signals often maintains reproductive boundaries between closely related, co‐occurring species. However, contrasting sources of selection, such as interspecific competition, can lead to weak divergence or even convergence of sexual signals in sympatry. When signals converge, assortative mating can be maintained if the mating preferences of females diverge in sympatry (reproductive character displacement; RCD), but there are few explicit examples. Pied flycatchers (Ficedula hypoleuca) are sympatric with collared flycatchers (F. albicollis) on the Baltic island of Öland, where males from both species compete over nestboxes, their songs converge, and the two species occasionally hybridize. We compare song discrimination of male and female pied flycatchers on Öland and in an allopatric population on the Swedish mainland. Using field choice trials, we show that male pied flycatchers respond similarly to the songs of both species in sympatry and allopatry, while female pied flycatchers express stronger discrimination against heterospecific songs in sympatry than in allopatry. These results are consistent with RCD of song discrimination of female pied flycatchers where they co‐occur with collared flycatchers, which should maintain species assortative mating despite convergence of male sexual signals.     

Song perception among incipient species as a mechanism for reproductive isolation

Many functions in behavioural processes of small passerines are regulated via vocal messages. Song plays an important role in the development of reproductive barriers and thus playback experiments can often be used for investigating the potential for reproductive isolation through behavioural mechanisms. Moltoni's warbler Sylvia (cantillans) moltonii is characterized by diagnostic vocalizations and a peculiar pattern of distribution, being parapatric and partly sympatric with the nominate Sylvia c. cantillans. With this work, we test whether these two closely related taxa react equally to their own song and to the song of the other taxon, shedding light on whether they perceive each other's songs as coming from the same species. We carried out 184 playback experiments within the mainland range of the two forms. We judged the response of the bird on a scale of scores. Each taxon responded more strongly to playback when faced with the song of its own taxon. This held true when applied only to males or females. Additionally, birds tested for both songs showed a stronger response to the song of their own taxon. The distributional context (sympatry vs. allopatry) did not affect bird response. Results indicate that a certain degree of reproductive isolation between the two taxa (because of diverged mate recognition systems) already exists; consistently with genetic data and with the peculiar pattern of distribution; this suggests that the two taxa have reached species status.