以鸟类视觉模型揭示中杜鹃对冠纹柳莺的卵色模拟(英文)

于2009年4—7月,采用光谱仪量化卵色和建立鸟类视觉模型的方法,在贵州宽阔水自然保护区对中杜鹃(Cuculus saturatus)寄生冠纹柳莺(Phylloscopus reguloides)的卵色模拟进行了研究。中杜鹃产白色卵带极少数而微小的棕色斑,明显大于宿主卵,重2.06g,体积1.91cm3。从人眼看,中杜鹃卵对宿主卵在很大程度上是模拟的,但视觉模型表明,两者的卵色在色调和色度上都完全分离,揭示了人眼探测不到的卵色模拟情况。该文首次对中杜鹃的雏鸟特征进行描述,在4日龄以后雏鸟嘴裂中出现三角形黑斑,并随着日龄的增长而更加明显,这种特征在霍氏中杜鹃(C.optatus)的雏鸟中也存在,但未见于其他种类的杜鹃雏鸟。     

中杜鹃在3种宿主巢中寄生繁殖

了解杜鹃对其宿主的选择和寄生情况,能为两者间的协同进化研究提供重要的基础资料。2012和2013年每年的4～8月,在贵州宽阔水国家级自然保护区对不同生境类型中的鸟巢进行搜索监测,记录到4例中杜鹃(Cuculus saturatus)寄生繁殖现象,其宿主分别是暗绿绣眼鸟(Zosterops japonicus)、棕腹柳莺(Phylloscopus subaffinis)和黄喉鹀(Emberiza elegans),其中棕腹柳莺和暗绿绣眼鸟是首次记录到被中杜鹃寄生。中杜鹃卵重(2.39±0.14)g(n=3),体积(2.24±0.18)cm3(n=3),通过T检验方法发现中杜鹃卵极显著大于暗绿绣眼鸟和棕腹柳莺卵(P0.001),同时在形状上也与两者极不相似,中杜鹃卵呈明显的长椭圆形,与黄喉鹀的卵在大小上无显著差异(P=0.1)。反射光谱的分析结果发现,寄生于不同宿主巢的中杜鹃卵在背景色和斑点上都具有一定差异,这表明中杜鹃的卵可能存在基因族群的分化。     

Geographic variation in parasitism rates of two sympatric cuckoo hosts in China

Rates of brood parasitism vary extensively among host species and populations of a single host species. In this study, we documented and compared parasitism rates of two sympatric hosts, the Oriental Reed Warbler (Acrocephalus orientalis) and the Reed Parrotbill (Paradoxornis heudei), in three populations in China. We found that the Common Cuckoo (Cuculus canorus) is the only parasite using both the Oriental Reed Warbler and Reed Parrotbill as hosts, with a parasitism rate of 22.4%-34.3% and 0%-4.6%, respectively. The multiple parasitism rates were positively correlated with local parasitism rates across three geographic populations of Oriental Reed Warbler, which implies that higher pressure of parasitism lead to higher multiple parasitism rate. Furthermore, only one phenotype of cuckoo eggs was found in the nests of these two host species. Our results lead to two conclusions: (1) The Oriental Reed Warbler should be considered the major host of Common Cuckoo in our study sites; and (2) obligate parasitism on Oriental Reed Warbler by Common Cuckoo is specialized but flexible to some extent, i.e., using Reed Parrotbill as a secondary host. Further studies focusing on egg recognition and rejection behaviour of these two host species should be conducted to test our predictions.     

大杜鹃和东方大苇莺卵的人工孵卵期和孵化率比较

大杜鹃(Cuculus canorus)是一种专性巢寄生鸟类,进化出了一系列适应对策,如雏鸟普遍出壳较早等,以更好适应寄生生活。本研究使用恒温自动孵化箱对25枚大杜鹃卵和20枚其宿主东方大苇莺(Acrocephalus orientalis)卵进行人工孵化,并对孵卵期的卵重进行连续测量。结果表明,在人工孵化条件下,大杜鹃卵的孵化率(76%)极显著高于东方大苇莺(40%)(χ~2=25.144,df=1,P0.01)。尽管大杜鹃的卵鲜重(t=7.447,df=43,P0.01)和卵体积(t=8.817,df=43,P0.01)均极显著大于东方大苇莺,但两种鸟卵的孵卵期不存在显著性差异(t=1.006,df=16,P0.05)。     

The evolution of host‐specific variation in cuckoo eggshell strength

Cuckoo eggs are renowned for their mimicry of different host species, leading to the evolution of host‐specific races (or ‘gentes’) defined by egg colour and pattern. This study aims to test the prediction that another property of parasitic eggs, namely shell strength, might also have experienced divergent selection within cuckoo species. Host races of the common cuckoo Cuculus canorus encountering stronger host rejection have thicker‐shelled eggs than those parasitising less discriminating species, as expected if egg strengthening discourages host rejection. Moreover, in the diederik cuckoo Chrysococcyx caprius, eggshell thickness was correlated across cuckoo gentes and host species, as expected if eggshell strength has been involved in coevolutionary interactions. This is the first report of host‐specific differences in cuckoo egg properties other than colour and pattern and lends correlational support to the hypothesis that the strong eggshells of brood parasites are an adaptation to reduce host rejection.     

Brood parasitism and egg matching in the Red-chested Cuckoo Cuculus solitarius in southern Africa

Host usage and relative rates of egg matching were investigated in the Red-chested Cuckoo Cuculus solitarius in southern Africa, using nest record cards and museum collections. Eighteen host species were found to be parasitized at varying degrees of intensity (0.14–12.5%). The most commonly recorded parasitized host, the Cape Robin Cossypha caffra , had a relatively low rate of parasitism (2.46%). The host species experiencing the most recorded pressure from parasitism was the Bearded Robin Erythropygia quadrivirgata , with 12.5% parasitism. Human perception of cuckoo/host egg matching was assessed for parasitized clutches of host species in museum egg collections. Eggs of three different cuckoo egg morphs were scored as matching those of the host species on a 1–5 scale. Perfect/good matching was recorded for eggs found in Chorister Cossypha dichroa , Heuglin's Cossypha heuglini and Natal Robins' Cossypha natalensis clutches. Poor and very poor matching was evident for cuckoo eggs found in four host species' clutches: the Cape Robin, Stonechat Saxicola torquata , Cape Rockthrush Monticola rupestris and Black Flycatcher Melaenornis pammelaina . Available evidence suggests that the Red-chested Cuckoo parasitizes hosts in a particular environment (low vegetation and trees). Good to intermediate matching was recorded with only 47% of host eggs, and with only 28.5% of Cape Robin clutches. A relatively high degree of host specificity, however, is suggested by the nest record card data, which indicate that species with large numbers of records are not those with the highest rates of parasitism.     

European Cuckoo Cuculus canorus parasitism and host's rejection behaviour in a heavily parasitized Great Reed Warbler Acrocephalus arundinaceus population

An unusually high frequency (64%) of European Cuckoo Cuculus canorus parasitism was found in Great Reed Warbler Acrocephalus arundinaceus clutches in central Hungary. Sixty-four per cent of the parasitized clutches contained one Cuckoo egg, 23% contained two, 10% had three and 3% had four. This means that 58% of the Cuckoo eggs were found in multiply parasitized clutches. In multiple parasitism the laying second Cuckoo removed an egg from the clutch randomly, so preferred neither the host eggs, nor the concurrent Cuckoo egg. Host response towards the parasitic eggs showed 66% acceptance, 12% ejection, 20% desertion and 2% egg burial. We found great variation in both the host and the parasitic egg colour and pattern. This reduces the chance that the parasitic egg's appearance matched that of the hosts' but, in spite of this, almost perfect mimesis was found in 28% of the Cuckoo eggs. Poorly mimetic Cuckoo eggs were more frequently rejected by Great Reed Warblers than parasite eggs that were very similar to the host eggs. This high level of mimicry sometimes makes it difficult for the observer to identify the parasitic egg, especially when it is similar in size to the host eggs. It is also difficult for the host, as shown by the relatively high recognition error and ejection cost.     

How to hatch from an egg of great structural strength. A study of the Common Cuckoo

Brood parasitism represents a unique mode of avian reproduction that requires a number of adaptations. For example, to reduce chances of puncture ejection of their eggs by small hosts, brood parasites may have been selected for laying eggs of unusually great structural strength. However, great structural strength of eggshells should hinder hatching. The goals of our study were to establish if chicks of the Common Cuckoo Cuculus canorus have more difficulty with hatching out of their strong eggs than chicks of species with eggs of similar size, and whether they possess any mechanisms facilitating hatching. To achieve these goals, we compared hatching pattern and selected body characteristics of chicks of the Common Cuckoo with those of another altricial species with eggs of a similar size, the Great Reed Warbler Acrocephalus arundinaceus . Although the rate of pecking was similar in the two species, the Common Cuckoo chicks started pecking earlier in relation to their emergence and consequently required more time and a greater cumulative number of pecks for breaking open their eggs than did young Great Reed Warblers. The two species also differed with respect to the pattern of opening their shells; in contrast to the warbler chicks, which enlarged the original pip circularly, the cuckoo chicks opened the egg by systematically creating a long narrow slit until they emerged. Finally, our study of hatched young revealed several differences; the Cuckoo hatchlings were significantly heavier, had a longer forearm, and their egg tooth was located significantly farther from the tip of the beak. The edge used for cutting through the shell was also significantly longer than that of hatchling Great Reed Warblers. To conclude, our data suggest that hatching is more difficult for a Cuckoo than for a Great Reed Warbler and that Cuckoos possess several mechanisms to overcome the problems of hatching from a structurally strong egg.     

Breeding success of common cuckoos Cuculus canorus parasitising four sympatric species of Acrocephalus warblers

We investigated the level of parasitism, egg mimicry and breeding success of cuckoos parasitising four sympatric species of Acrocephalus warblers in southern Moravia, Czech Republic. The parasitism rate was highest in the marsh warbler Acrocephalus palustris (44.8%) followed by great reed warbler A. arundinaceus (33.8%), sedge warbler A. schoenobaenus (26.5%) and reed warbler A. scirpaceus (11.6%). Although the cuckoo eggs showed a high level of mimicry the eggs of the marsh warbler this host species rejected 72% of the cuckoo eggs, resulting in a cuckoo breeding success of only 4.3%. Cuckoo eggs laid in great reed warbler and reed warbler nests showed a similar hatching success, but the cuckoo chicks survived better in great reed warbler nests, resulting in a breeding success of 30.4%, as compared to 16.4% in nests of the reed warbler. The relationship between the level of parasitism, host rejection of cuckoo eggs, cuckoo chick survival and breeding success is discussed for the four host species.     

Egg colour matching in an African cuckoo, as revealed by ultraviolet-visible reflectance spectrophotometry

Despite major differences between human and avian colour vision, previous studies of cuckoo egg mimicry have used human colour vision (or standards based thereon) to assess colour matching. Using ultraviolet-visible reflectance spectrophotometry (300-700 nm), we measured museum collections of eggs of the red-chested cuckoo and its hosts. The first three principal components explained more than 99% of the variance in spectra, and measures of cuckoo host egg similarity derived from these transformations were compared with measures of cuckoo host egg similarity estimated by human observers unaware of the hypotheses we were testing. Monte Carlo methods were used to simulate laying of cuckoo eggs at random in nests. Results showed that host and cuckoo eggs were very highly matched for an ultraviolet versus greenness component, which was not detected by humans. Furthermore, whereas cuckoo and host were dissimilar in achromatic brightness, humans did not detect this difference. Our study thus reveals aspects of cuckoo-host egg colour matching which have hitherto not been described. These results suggest subtleties and complexities in the evolution of host-cuckoo egg mimicry that were not previously suspected. Our results also have the potential to explain the longstanding paradox that some host species accept cuckoo eggs that are non-mimetic to the human eye.     

Climate change and coevolution in the cuckoo–reed warbler system

The evolution of traits in hosts may be influenced by their parasites and vice versa and a coevolutionary arms race often develops between the two. As part of such an arms race, the common cuckoo mimics the eggs of its hosts to avoid egg rejection. Traits related to this arms race may also be influenced by climatic conditions, such as temperature, affecting, for example, food availability and, thus, female condition and egg size (therefore may reflect Bergmann’s rule or the resource rule). The potential interaction between coevolution and climate has rarely been studied. We investigated whether egg and body size of cuckoos and reed warblers from Britain and Denmark had undergone change between 1868 and 1956, and whether such changes were correlated with climatic factors. Cuckoo egg size decreased during the studied period while warbler egg size remained stable. Hence, cuckoo and warbler eggs have become more similar in size over time. Cuckoo egg volume decreased with increasing annual precipitation, but annual precipitation decreased over time. Warbler egg volume increased with spring temperatures (which could not reflect Bergmann’s rule, but may support the resource rule). Hence, it seems that the measured climatic indices did not affect cuckoo egg size but may in part affect warbler egg size. Therefore, the decrease in cuckoo egg size may be the result of the coevolutionary arms race. Body and egg sizes in the cuckoos were negatively correlated whereas warbler body and egg sizes were uncorrelated, suggesting that selection probably acted on egg size directly and not via selection on body size. Taken together, these findings may indicate that climate change, the coevolutionary arms race, or both, affected egg sizes. It is suggested that drawing conclusions regarding the arms race without taking into account other selective pressures (e.g., climate) may confound conclusions regarding parasite-host systems.     

Egg shape mimicry in parasitic cuckoos

Parasitic cuckoos lay their eggs in nests of host species. Rejection of cuckoo eggs by hosts has led to the evolution of egg mimicry by cuckoos, whereby their eggs mimic the colour and pattern of their host eggs to avoid egg recognition and rejection. There is also evidence of mimicry in egg size in some cuckoo–host systems, but currently it is unknown whether cuckoos can also mimic the egg shape of their hosts. In this study, we test whether there is evidence of mimicry in egg form (shape and size) in three species of Australian cuckoos: the fan‐tailed cuckoo Cacomantis flabelliformis, which exploits dome nesting hosts, the brush cuckoo Cacomantis variolosus, which exploits both dome and cup nesting hosts, and the pallid cuckoo Cuculus pallidus, which exploits cup nesting hosts. We found evidence of size mimicry and, for the first time, evidence of egg shape mimicry in two Australian cuckoo species (pallid cuckoo and brush cuckoo). Moreover, cuckoo–host egg similarity was higher for hosts with open nests than for hosts with closed nests. This finding fits well with theory, as it has been suggested that hosts with closed nests have more difficulty recognizing parasitic eggs than open nests, have lower rejection rates and thus exert lower selection for mimicry in cuckoos. This is the first evidence of mimicry in egg shape in a cuckoo–host system, suggesting that mimicry at different levels (size, shape, colour pattern) is evolving in concert. We also confirm the existence of egg size mimicry in cuckoo–host systems.     

Responses of the Bush Warbler Cettia diphone to artificial eggs of Cuculus cuckoos in Japan

Artificial eggs of six different colours and control eggs of Bush Warblers Cettia diphone were introduced into nests of Bush Warblers, a host of both Little and Himalayan Cuckoos Cuculus poliocephalus and C. saturatw in Japan. All control (chocolate-brown) and artificial red eggs were accepted; all grey and all white eggs were rejected. The rejection rates of orange, pink and orange spots on grey eggs were 8%, 369; and 55%, respectively. Bush arblers are more likely to reject eggs the more dissimilar they are from their own. The results strengthen the possibility that the chocolate-brown eggs of Little and Himalayan Cuckoos may have evolved through the discriminative ability of Bush Warblers and their intolerance towards dissimilar eggs.     

八声杜鹃在两种缝叶莺巢中寄生繁殖

了解杜鹃对不同寄主的选择和利用,可为研究两者之间的协同进化提供重要基础资料。2012至2014年每年的3~6月,在广西弄岗地区共记录到3例八声杜鹃(Cacomantis merulinus)的寄生繁殖,其中,2例寄生于栗头缝叶莺(Orthotomus cuculatus)的巢(寄生率1.4%,n=142),其中2枚杜鹃卵均为白色具棕色斑点;1例寄生于黑喉缝叶莺(O.atrogularis)(寄生率10%,n=10),杜鹃卵为白色具棕色斑点。栗头缝叶莺为八声杜鹃寄主的首次报道,而两种缝叶莺均首次在中国记录到被八声杜鹃寄生。八声杜鹃的卵重为(1.45±0.09)g,卵体积为(1.46±0.07 cm3)(n=3),其卵大小和重量均显著大于栗头缝叶莺和黑喉缝叶莺的卵(t检验,P0.001)。     

Rapid increase in cuckoo egg matching in a recently parasitized reed warbler population

Parasitic cuckoos lay eggs that mimic those of their hosts, and such close phenotypic matching may arise from coevolutionary interactions between parasite and host. However, cuckoos may also explicitly choose hosts in a way that increases degree of matching between eggs of cuckoos and parasites, with female preference for specific host phenotypes increasing the degree of matching. We tested for temporal change in degree of matching between eggs of the parasitic European cuckoo (Cuculus canorus) and its reed warbler (Acrocephalus scirpaceus) host during 24 consecutive years in a recently parasitized reed warbler population. Cuckoo-host egg matching in an ultraviolet-brownness component yielding most of the chromatic variance of eggs improved during the study period. Improved matching was not due to changes in cuckoo egg phenotype. Cuckoo eggs matched host eggs for ultraviolet-brownness within nests irrespective of duration of sympatry. Ultraviolet-brownness of cuckoo eggs was similar to that of reed warbler eggs at parasitized nests, but differed from that of reed warbler eggs at unparasitized nests. These findings provide tentative support for the cuckoo preference hypothesis suggesting that cuckoo-host egg matching could partially be due to cuckoo females selecting host nests based on the appearance of their eggs.     

小杜鹃对强脚树莺的巢寄生及其卵色模拟

于1999-2009年的鸟类繁殖季(4-8月)对贵州宽阔水自然保护区内的强脚树莺(Cettia fortipes)的繁殖进行监测,并采用光纤光谱仪,通过主成分分析、反射光谱、罗宾逊投射等方法对强脚树莺的卵色与小杜鹃(Cuculus poliocephalus)对其寄生的卵色模拟程度进行分析。研究结果表明,强脚树莺的繁殖成效在不同年份间均无显著差异,但其巢被捕食率和巢被寄生率都较高,分别为49.26%和9.18%。通过反射光谱分析,表明小杜鹃卵在色调和色度上高度模拟强脚树莺的卵,但其亮度高于宿主卵,而且在人眼无法探测的紫外光部分存在差异。     

The common cuckoo Cuculus canorus is not locally adapted to its reed warbler Acrocephalus scirpaceus host

The obligate avian brood parasitic common cuckoo Cuculus canorus comprises different strains of females that specialize on particular host species by laying eggs of a constant type that often mimics those of the host. Whether cuckoos are locally adapted for mimicking populations of the hosts on which they are specialized has never been investigated. In this study, we first explored the possibility of local adaptation in cuckoo egg mimicry over a geographical mosaic of selection exerted by one of its main European hosts, the reed warbler Acrocephalus scirpaceus. Secondly, we investigated whether cuckoos inhabiting reed warbler populations with a broad number of alternative suitable hosts at hand were less locally adapted. Cuckoo eggs showed different degrees of mimicry to different reed warbler populations. However, cuckoo eggs did not match the egg phenotypes of their local host population better than eggs of other host populations, indicating that cuckoos were not locally adapted for mimicry on reed warblers. Interestingly, cuckoos exploiting reed warblers in populations with a relatively larger number of co-occurring cuckoo gentes showed lower than average levels of local adaptation in egg volume. Our results suggest that cuckoo local adaptation might be prevented when different cuckoo populations exploit more or fewer different host species, with gene flow or frequent host switches breaking down local adaptation where many host races co-occur.     

SOME NOTES ON THE BREEDING BIOLOGY OF THE STRIPED CUCKOO

Steyn, P. 1973. Some notes on the breeding biology of the Striped Cuckoo. Ostrich 44: 163–169.

Information is presented on the breeding biology of the Striped Cuckoo, a species for which little authentic material exists. A number of cases of parasitism of the Arrow-marked Babbler are given. Pre-laying behaviour is similar to that of the Jacobin Cuckoo. The blue egg of the cuckoo may be distinguished on several minor points, but mainly because it is rounder and broader than those of the host species. The growth and development of a nestling is outlined up to its ninth day when it was killed by a snake. It was reared to this stage with three babbler chicks, probably because several babblers contribute to feeding the nestlings. The cuckoo gains weight very rapidly, and it is suggested that this is because of its brighter gape and more intense gaping response which ensure that it is fed preferentially. Anti-predator devices such as open-gaped lunges, jerking movements of the body and the exudation of a vile-smelling brown fluid are described. The nestling cuckoo's call is identical to that of the babblers. The juvenal may be fed by its foster parents For at least 36 days after leaving the nest.     

Continuous Variation Rather than Specialization in the Egg Phenotypes of Cuckoos (Cuculus canorus) Parasitizing Two Sympatric Reed Warbler Species

The evolution of brood parasitism has long attracted considerable attention among behavioural ecologists, especially in the common cuckoo system. Common cuckoos (Cuculus canorus) are obligatory brood parasites, laying eggs in nests of passerines and specializing on specific host species. Specialized races of cuckoos are genetically distinct. Often in a given area, cuckoos encounter multiple hosts showing substantial variation in egg morphology. Exploiting different hosts should lead to egg-phenotype specialization in cuckoos to match egg phenotypes of the hosts. Here we test this assumption using a wild population of two sympatrically occurring host species: the great reed warbler (Acrocephalus arundinaceus) and reed warbler (A. scirpaceus). Using colour spectrophotometry, egg shell dynamometry and egg size measurements, we studied egg morphologies of cuckoos parasitizing these two hosts. In spite of observing clear differences between host egg phenotypes, we found no clear differences in cuckoo egg morphologies. Interestingly, although chromatically cuckoo eggs were more similar to reed warbler eggs, after taking into account achromatic differences, cuckoo eggs seemed to be equally similar to both host species. We hypothesize that such pattern may represent an initial stage of an averaging strategy of cuckoos, that – instead of specializing for specific hosts or exploiting only one host – adapt to multiple hosts.     

Coevolution of an avian host and its parasitic cuckoo

Abstract We use a quantitative genetic model to examine the coevolution of host and cuckoo egg characters (termed "size" as a proxy for general appearance), host discrimination, and host and cuckoo population dynamics. A host decides whether to discard an egg using a comparison of the sizes of the eggs in her nest, which changes as host and cuckoo eggs evolve. Specifically, we assume that the probability that she discards the largest egg in her nest depends on how much larger it is than the second largest egg. This decision rule (i.e., the acceptable difference in egg sizes) also evolves, changing both the chance of successful rejection of a cuckoo egg in parasitized nests and the chance of mistaken rejection of a host egg in both parasitized and unparasitized nests. We find a stable equilibrium for coexistence of the host and cuckoo where there is cuckoo egg mimicry, evolutionary displacement of the host egg away from the cuckoo egg phenotype, and host discrimination against unusual eggs. Both host discrimination and host egg displacement are fairly weak at the equilibrium. Cuckoo egg mimicry, although imperfect, usually evolves more extensively and quickly than the responses of the host. Our model provides evidence for both the evolutionary equilibrium and evolutionary lag hypotheses of host acceptance of parasitic eggs.