Call divergence is correlated with geographic and genetic distance in greenish warblers (Phylloscopus trochiloides): a strong role for stochasticity in signal evolution?

Divergence in signalling systems might play a central role in speciation. To assess the importance of possible causes of signal divergence, we examine two types of vocalizations within a geographically variable species complex, the greenish warblers (Phylloscopus trochiloides Sundevall). Calls, which are used by both sexes throughout the year, and songs, which are sung primarily by breeding males, differ distinctly between two distinct Siberian forms. Through a ring of southern populations that connect the northern forms, signal divergence is correlated with both geographic distance and genetic divergence. Calls and songs differ in their particular patterns of geographic variation, probably because of the larger influence of sexual selection on songs than on calls. These patterns are supportive of neither acoustic adaptation nor morphology being major drivers of divergence in vocalizations. Rather, these results support the importance of stochastic evolution of communication systems in the evolution of new species.     

云南柳莺鸣声特点及地理差异的初步分析

1997、2002和2005年5～7月,在陕西省、四川省和甘肃省内共7个地区采集云南柳莺(Phylloscopus yunnanensis)的鸣声样本,并分析采自不同地区的共40只个体的鸣声.发现云南柳莺的鸣唱(song)和鸣叫(calls)在同一地区存在一定的差异.比较99段云南柳莺的鸣唱声,发现持续时间与间隔时间之间存在显著的负相关Pearson,r=-0.276,P=0.006,t=99).单因素方差分析表明,不同地区云南柳莺鸣唱的最高频率、最低频率和持续时间没有显著差异,而音节的间隔时间及音素数量均存在显著差异(P=0.006;P=0.000).结果表明,不同海拔、地区之间个体鸣唱音节间隔时间与海拔成正相关(Pearson,r=0.425,P=0.03,n=26).     

Ring species – Do they exist in birds?

According to current theory, the splitting of a single species into two is best observed by a ring of intergrading populations which occupy a ring-shaped distribution area and whose terminal populations not only meet but overlap and co-occur without or with only little hybridization. The three most discussed examples in birds are revisited here. The great tit complex (Parus major s. l.) turned out to be an assemblage of four subspecies groups forming a secondary ring of population. The herring gull/lesser black-backed gull complex (Larus argentatus s. l.) forms a circumpolar circle of intergrading populations, but lacks the crucial cornerstone, the geographical overlap. The greenish warbler complex (Phylloscopus trochiloides s. l.) is close to the ideal ring species, but the ring is not complete and the Siberian zone of overlap needs further clarification and characterisation.     

Lack of habitat segregation and no interspecific territoriality in three syntopic cryptic species of the golden‐spectacled warblers Phylloscopus (Seicercus) burkii complex

It is widely accepted that exploitative competition prevents the coexistence of any two or more closely‐related species unless differences exists in their ecological niches and resource use. In sibling bird species, exploitative competition is reduced mainly by spatial segregation of competing species. Spatial segregation can be achieved in two basic ways: by using different habitats or microhabitats that each species is predominately exploiting, or by interspecific territoriality. To our knowledge, either habitat segregation or interspecific territoriality or both have been found in all dyads and groups of sympartic sibling bird species studied so far. In this study, we investigated the coexistence of three cryptic sibling species of the golden‐spectacled warblers Phylloscopus burkii complex in Hunan Province, China: Ph. tephrocephalus, Ph. omeiensis and Ph. valentini. We analyzed their habitat preferences, and spatial interrelations between the species. In order to analyze either presence or absence of interspecific territoriality, we additionally performed con‐ and heterospecific playback experiments. Contrary to expectations, we found no evidences for either habitat segregation or interspecific territoriality in these three species.     

河北塞罕坝褐柳莺繁殖期鸣唱变化

2017年5至8月,在河北省塞罕坝录制了80只繁殖期雄性褐柳莺(Phylloscopusfuscatus)的鸣唱,对其鸣唱句型和句子参数以及音节类型的变化进行了定量分析。褐柳莺鸣唱的两种主要句型——S-song和V-song,前者(F_(3,75)=17.983,P 0.05)和后者(F_(3,75)=17.983,P 0.05)的使用频次在不同月份间均存在显著差异。S-song的使用频次多于V-song的使用频次,且二者在繁殖季的变化表现出相反的趋势:S-song使用频次在5月和8月较高,在6月最低;而V-song的使用频次却是在6月达到最高值,在5月和8月较低。不同月份鸣唱句子最高频率(F_(3,75)=4.841,P 0.05)、句子频宽(F_(3,75)=3.814,P 0.05)和句子峰频率(F_(3,75)=3.793,P 0.05)均存在显著差异。通过80只个体的2 400个鸣唱语句得到了65种音节类型,其中,80%以上的音节类型在每个月中均有分布,但每月音节类型数存在显著差异(F_(3,75)=5.456,P 0.05)。音节类型数量随繁殖季的变化表现为,7月褐柳莺鸣唱中具有最多的音节类型数(19.79±3.69),5月和8月音节类型数较少,8月最少。     

Does acoustically simulated predation risk affect settlement and reproduction of a migratory passerine?

Nest predation is one of the most important drivers of avian life history evolution and population dynamics. Increasing evidence suggests that birds are able to assess nest predation risk and avoid settling in high‐risk areas to increase their reproductive performance. However, the cues used for settlement decisions are poorly known in most species. Population sizes of the migratory wood warbler Phylloscopus sibilatrix are characterized by strong annual fluctuations, which are negatively correlated with the number of forest rodents. Wood warblers might avoid rodent‐rich areas to reduce predation risk arising either from rodents, from rodent‐hunting predators attracted to such areas or from predators not linked to rodents. To evaluate these hypotheses, we conducted a large‐scale field experiment to test whether wood warblers avoided settling in plots with high predation risk simulated by broadcasting vocalizations of rodents or predators. Moreover, we tested whether reproductive performance varied in relation to simulated predation risk. Settlement patterns did not differ between plots with rodent, predator and noise control treatments. Likewise, measures of reproductive performance did not seem to differ across treatments. Thus, the broadcasted vocalizations of rodents and predators did not seem to be perceived as threat by wood warblers. Alternatively, the species might use other cues than those presented here, either other acoustic cues, visual and/or olfactory cues or a combination of cue types during settlement. Further experimental investigations to pin point cues and senses relevant for settlement decisions in wood warblers and birds in general are needed to better understand their life history and population dynamics.     

Recurrent selection explains parallel evolution of genomic regions of high relative but low absolute differentiation in a ring species

Recent technological developments allow investigation of the repeatability of evolution at the genomic level. Such investigation is particularly powerful when applied to a ring species, in which spatial variation represents changes during the evolution of two species from one. We examined genomic variation among three subspecies of the greenish warbler ring species, using genotypes at 13 013 950 nucleotide sites along a new greenish warbler consensus genome assembly. Genomic regions of low within‐group variation are remarkably consistent between the three populations. These regions show high relative differentiation but low absolute differentiation between populations. Comparisons with outgroup species show the locations of these peaks of relative differentiation are not well explained by phylogenetically conserved variation in recombination rates or selection. These patterns are consistent with a model in which selection in an ancestral form has reduced variation at some parts of the genome, and those same regions experience recurrent selection that subsequently reduces variation within each subspecies. The degree of heterogeneity in nucleotide diversity is greater than explained by models of background selection, but is consistent with selective sweeps. Given the evidence that greenish warblers have had both population differentiation for a long period of time and periods of gene flow between those populations, we propose that some genomic regions underwent selective sweeps over a broad geographic area followed by within‐population selection‐induced reductions in variation. An important implication of this ‘sweep‐before‐differentiation’ model is that genomic regions of high relative differentiation may have moved among populations more recently than other genomic regions.     

Laying dates and clutch size of open-nesting passerines in the Czech Republic: a comparison of systematically and incidentally collected data

Data are presented on laying date and clutch size (n) of Blackcap Sylvia atricapilla (1434), Song Thrush Turdus philomelos (298), Garden Warbler S. borin (260), Yellowhammer Emberiza citrinella (254), Blackbird T. merula (231), Chaffinch Fringilla coelebs (206), Chiffchaff Phylloscopus collybita (202), Icterine Warbler Hippolais icterina (95) and Dunnock Prunella modularis (89), recorded during an intensive study (since 1993) in woodland habitats of Eastern Bohemia, Czech Republic (49 °54 ′N, 16 °02 ′E). Results are compared with two other data sets (a local study and national nest record scheme) collected by Czech volunteer ornithologists during 1945–85 and with other published data from central Europe. There is no evidence that the laying dates and clutch size of the nine species changed at the regional scale over the past decades, and mean values generally match their known geographical variability. Significant species differences were found among the three Czech data sets in both laying dates and clutch size. Most of the differences could be explained as an artefact of seasonally biased sampling of nests, which in turn influenced mean clutch size. Even a systematic study with a standard searching effort on fixed plots provided biased data for some species.     

Positive correlations between pre‐ and post‐copulatory sexual traits in warblers

Theoretical models predict that investment in pre‐copulatory and post‐copulatory sexually selected traits should trade‐off. At the macroevolutionary scale, the majority of studies to date have focused on male weaponry as the target of pre‐copulatory sexual selection, but the trade‐off should equally apply to traits used to attract females, such as bird song and plumage. We studied the Old World leaf warblers (Phylloscopidae), a group of socially monogamous songbirds that experience relatively high levels of sperm competition. We examined the relationships between song duration and number of elements in the song with sperm length across 21 species, and between the same song variables and combined testes mass in a subset of these species (n = 10). Across species, these song variables and testes mass/sperm length were generally positively correlated, albeit not statistically significantly so or with borderline significance. In contrast to theory, we found no evidence for negative associations between pre‐ and post‐copulatory traits. We argue that this is a consequence of males of some species investing more into overall fertilization success (i.e. the sum of pre‐ and post‐copulatory sexual selection) than males of other species, and high fertilization success is achieved through investment into both mate attraction and sperm competition.