An example of character release in host selection and egg colour of cuckoos Cuculus spp. in Japan

We have studied in northern Japan the hitherto unidentified eggs of a species of cuckoo in the nests of the Bush Warbler Cettia diphone . The cuckoo in question appeared to be the Himalayan Cuckoo Cuculus saturatus which parasitizes mainly the Willow Warbler Phylloscopus occipitalis in southern Japan. The egg colour in this northern Cuckoo was chocolate-brown or orange-brown, similar to that of the Bush Warbler but unlike that of the southern Himalayan Cuckoo. Egg size was significantly larger than that of the southern Himalayan Cuckoo and instead similar to that of the Little Cuckoo C. poliocephalus which uses the same host species in southern Japan, to which the Little Cuckoo is confined. The shift in host species and egg colour in the northern Himalayan Cuckoo seems to be a case of character release in the absence of the Little Cuckoo.     

Brood-parasitism by the Shining Cuckoo Chrysococcyx lucidus at Kaikoura, New Zealand

For three seasons starting in 1976 I studied the breeding of Shining Cuckoos Chrysococcyx lucidus in forest near Kaikoura, New Zealand. There is no evidence that the cuckoo parasitizes any host on mainland New Zealand other than the Grey Warbler Gerygone igata. A nestling cuckoo returned to within 1 km of its natal site in a subsequent breeding season, presumably after migrating beyond New Zealand. Empirical and theoretical estimates of the area occupied by Shining Cuckoos while breeding are given. Cuckoos near Kaikoura laid during ten weeks, the modal week of laying following seven weeks after the presumed peak of arrival of birds in New Zealand. First clutches of the host escaped parasitism because they were laid before most cuckoos arrived. Parasitized clutches received one cuckoo egg which replaced a host's egg. It was laid before, just after or long after the host began incubating, and mimicry was lacking. Cuckoo eggs, which were about 8% of the adult cuckoo's weight, hatched in 14–17 days. The frequency of parasitism near Kaikoura was 55% of late clutches (n = 40).
At 3–7 days old, nestling Shining Cuckoos evicted from the nest all other contents. The nestling period was at least 19 days. Growth in weight followed a logistic curve and the equation is given. Just over half the cuckoo eggs produced fledglings. The effect of brood-parasitism on the Grey Warbler's productivity was small. Only 17% of late warbler eggs, and late eggs only, were prevented by parasitism from yielding fledglings. Late laying by some Shining Cuckoos (relative to the host's incubational cycle), and late eviction, often led to brief inter-specific competition among nestlings for food. The brief coexistence of young Warblers and Cuckoos in the nest may explain the apparent mimicry by newly-hatched Shining Cuckoos of the host's young.     

Abundance,feeding, and morphology of passerine birds at Kowhai Bush,Kaikoura, New Zealand

Abstract

Five-minute stationary counts of birds at Kowhai Bush over 17 months suggest that the scores for grey warblers and chaffinches may reflect vocal conspicuousness, rather than abundance, and that the scores for shining cuckoos, goldfinches, and redpolls may reflect vocal and visual conspicuousness in combination. Counts in three types of kanuka forest show that cuckoos, robins, and bellbirds favoured the more dense and diverse suocessional stages; that riflemen, brown creepers, fantails, chaffinches, goldfinches, and redpolls were most abundant in less mature habitats; and that warblers and silvereyes were almost uniformly common. Cuckoos and robins overlapped the most in use of habitat, robins and redpolls the least. For both warblers and robins, indices of abundance varied between two of the habitats in proportion to the densities of resident adults, permitting calibration of the indices. Excepting creepers and robins, native species were apparently less abundant at Kowhai Bush than in climax forest near Reefton. At Kowhai Bush in winter, creepers, warblers, and silvereyes (three of the four small native gleaners of foliage) collected prey almost entirely from kanuka, the dominant tree. Warblers foraged on 80% of occasions from living foliage, whereas creepers fed almost equally from trunks, branches, twigs, and leaves, and silvereyes concentrated on leaves and trunks. Creepers and silvereyes overlapped the most in use of feeding stations. Also, they were exclusively gleaners, whereas warblers caught prey on 40% of occasions by hovering. Warblers gleaned only in the upright position, but creepers and silvereyes often gleaned from vertical surfaces or by hanging upside down. The greatest overlap in feeding behaviour was between creepers and silvereyes. Data for four of the five small native insectivores show that warblers were half as heavy as creepers and silvereyes, and lighter on average than fantails. The tail was longer than the wing in fantails, shorter than the wing in silvereyes, and equal to the wing in warblers and creepers. The ratio of wing length to tarsometatarsus length was greatest for fantails (3.4), as befits an aerial feeder. Warblers, silvereyes, and fantails had bills that were wider than deep; the creeper’s was slightly deeper than wide. Silvereyes had the longest bill, and creepers the longest tarsometatarsus. Indices of morphological difference show that silvereyes and creepers differed least.     

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.     

Responses of breeding Reed Warblers Acrocephalus scirpaceus to mounts of Sparrowhawk Accipiter nisus, Cuckoo Cuculus canorus and Jay Garrulus glandarius

Mounts of a Cuckoo Cuculus canorus , a Sparrowhawk Accipiter nisus and a Jay Garrulus glandarius were presented at nests of Reed Warblers Acrocephalus scirpaceus at all breeding stages. Response strength could best be classified according to approach distance (to the mount) and vocalizations used. Warblers reacted much more aggressively to a mount on the nest than to one 3 m adjacent; they approached much more closely, were less likely to sing but often gave a frenzied-sounding rasp (which was never given to an adjacent mount). Reaction was more aggressive to a Cuckoo, which was often attacked, than to a Sparrowhawk, of which they were wary; response to a Jay was intermediate. Reaction was more aggressive after the clutch had been completed than before. After the brood had fledged, parents still responded strongly to the Jay and Sparrowhawk, but responded to the Cuckoo only briefly before ignoring it. These results are consistent with the Reproductive Value-Stimulus Value hypothesis. They also demonstrate that Reed Warblers can tell Cuckoos from Sparrowhawks, as can many non-host species. The close resemblance of these two species is often suggested to be mimicry of the Sparrowhawk by the Cuckoo but, if this is so, it is not successful. Alternatively, the similarity could be mimicry to reduce predation on the Cuckoos themselves, a chance resemblance, or due to similar selective forces acting on both species, as each would presumably benefit from reduced conspicuousness.     

Nest Sanitation Plays a Role in Egg Burial by Yellow Warblers

Some hosts of the brown-headed cowbird ( Molothrus ater ) possess defences that eliminate all or most parasitism costs. Yellow warblers ( Dendroica petechia ) bury cowbird eggs, possibly to clean nests rather than serving strictly as an anti-parasite defence, as non-egg-shaped objects have been ejected, buried, or deserted by other hosts. With two experiments, we tested the 'nest sanitation' hypothesis by recording warblers' responses to objects (1) similar in volume, mass, and colour to cowbird eggs, and (2) half the mass and volume (more easily ejected), placed into nests before and during incubation. We compared outcomes at control nests with responses to objects that were dissimilar (stars) and moderately similar (dumbbells) to eggs, and to real cowbird and warbler eggs. We tested whether rejection (1) declines from stars through dumbbells and real eggs, (2) is similar between stages, and (3) non-egg-shaped objects are ejected because this is the least costly rejection method. Large stars were rejected (most buried) significantly more frequently (43.8%) than cowbird eggs (16.3%) in pre-incubation, suggesting that warblers reject objects shaped unlike their own eggs to rid nests of debris. Objects spent less time in nests the more they diverged from eggs. Warblers rarely rejected large stars and dumbbells, and cowbird eggs during incubation, possibly because burial and desertion are too costly by this time. Responses to small stars and dumbbells, and to foreign yellow warbler eggs did not differ between stages; also warblers rejected stars, mostly by ejection and selective burial, more frequently (28.8%) than dumbbells (1.3%) and warbler eggs (0%). Rejection by yellow warblers, especially burial, may keep nests clean, but also functions in rejecting cowbird eggs.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

Common Cuckoo Cuculus canorus and host behaviour at Reed Warbler Acrocephalus scirpaceus nests

This study is based on continuous video recordings made at 53 Reed Warbler Acrocephalus scirpaceus nests each day during the laying period. Egg-laying by the Common Cuckoo Cuculus canorus was recorded in 14 (26.4%) of these nests. By analysing the activity of the host birds around and at the nest, we found that this is probably not the only cue used by the Common Cuckoo when locating suitable nests to parasitize. Furthermore, in most cases there was no significant difference between the length of time the host birds spent at the nest in the morning and afternoon, thus providing little support for the hypothesis that the Common Cuckoo lays in the afternoon because it is less likely to be seen by the nest owners then. Parasitized Reed Warblers rejected the Common Cuckoo egg more frequently when they observed the parasite at their nests. However, contrary to what should be expected, most Common Cuckoos laid their eggs in the presence of the host(s), and in general their egg-laying behaviour (for example duration of stay at the nest) was less secretive than described earlier. When partially depredating host clutches, Cuckoos showed the same behavioural pattern at parasitized and unparasitized nests, indicating that the latter may act as a potential reserve for egg-laying.     

Egg rejection behavior in a population exposed to parasitism: Village Weavers on Hispaniola

In contrast to African Village Weavers (Ploceus cucullatus)that are parasitized by Diederik Cuckoos (Chrysococcyx caprius),introduced weavers on Hispaniola existed without parasitismfor at least 2 centuries until the arrival of the Shiny Cowbird(Molothrus bonariensis) in the 1970s. Cruz and Wiley (1989)found that Hispaniolan weavers had a lower rejection rate offoreign eggs than African populations. Subsequently, Robertand Sorci (1999) and Lahti (2005, 2006) found that acceptanceof dissimilar eggs is not characteristic of the species throughoutits Hispaniolan range. In 1999–2002, we studied egg rejectionin Hispaniolan weavers on a broad regional scale. Rejectionincreased as experimental eggs became increasingly differentfrom the host eggs. Rejection rates for mimetic eggs, differentcolor eggs, different-spotting eggs, and cowbird eggs was 23.2%,33.3%, 61.5%, and 85.3%, respectively, with higher rejectionof cowbird eggs in areas where cowbirds were observed. Althoughrejection is likely to have a genetic component, the differencescould be due to phenotypic plasticity. Plasticity in egg rejectionmay be expected, given the potential cost of rejection and thespatiotemporal distribution of cowbirds. Thus, egg rejectionhas not necessarily decreased in Hispaniolan weavers, but itmay act in a plastic manner, increasing where cowbirds are present.     

Breeding ground correlates of the distribution and decline of the Common Cuckoo Cuculus canorus at two spatial scales

Many migratory bird species are undergoing population declines as a result of potentially multiple, interacting mechanisms. Understanding the environmental associations of spatial variation in population change can help tease out the likely mechanisms involved. Common Cuckoo Cuculus canorus populations have declined by 69% in England but increased by 33% in Scotland. The declines have mainly occurred in lowland agricultural landscapes, but their mechanisms are unknown. At both the local scale within the county of Devon (SE England) and at the national (UK) scale, we analysed the breeding season distribution of Cuckoos in relation to habitat variation, the abundance of host species and the abundance of moth species whose caterpillars are a key food of adult Cuckoos. At the local scale, we found that Cuckoos were more likely to be detected in areas with more semi‐natural habitat, more Meadow Pipits Anthus pratensis (but fewer Dunnocks Prunella modularis) and where, later in the summer, higher numbers of moths were captured whose larvae are Cuckoo prey. Nationally, Cuckoos have become more associated with upland heath characterized by the presence of Meadow Pipit hosts, and with wetland habitats occupied by Eurasian Reed Warbler Acrocephalus scirpaceus hosts. The core distribution of Cuckoos has shifted from south to north within the UK. By the end of 2009, the abundance of macro‐moth species identified as prey had also declined four times faster than that of species not known to be taken by Cuckoos. The abundance of these moths has shown the sharpest declines in grassland, arable and woodland habitats and has increased in semi‐natural habitats (heaths and rough grassland). Our study suggests that Cuckoos are likely to remain a very scarce bird in lowland agricultural landscapes without large‐scale changes in agricultural practices.     

BREEDING OF THE GREY WARBLER GERYGONE IGATA AT KAIKOURA, NEW ZEALAND

I studied the breeding of Grey Warblers Gerygone igata (Muscicapidae: Acanthizinae) in forest near Kaikoura, New Zealand, between 1976 and 1979. Only males sang and singing occurred all year. From late July to January pairs defended self-contained territories of 0·25–1·73 ha but they occupied larger home ranges when not breeding. Territorial adults were strictly sedentary all year. The average annual mortality of breeding adults was 18·5% and the predicted life-expectancy 4·9 years, which is remarkable in a bird weighing 6–7 g. The breeding season from first building to last fledging was six months long and it began early. Exceptionally, Grey Warblers may build and lay before the shortest day. As the season progressed warblers nested lower on average, both in absolute terms and relative to the tree nested in and canopy at the site. Warblers built in 7–27 days then delayed up to eight days before laying. Only females built and at no stage of breeding did males feed their mates. Both sexes fed the young. Grey Warblers laid for 15–16 weeks of the year and first clutches were laid asynchronously during 5–6 weeks. Eggs of a clutch appeared at two-day intervals and each egg weighed 1·5 g when fresh (23% of mean adult weight). Clutch size was nearly constant (mean 3·9, mode 4, range 3–5). The incubation period was 17–21 days (mean 19·5 days) and the nestling period 15–19 days (mean 17·2 days). On average the clutch hatched over 1·4 days, even though incubation commenced with the laying of the last egg. Nestlings reached maximum weight on Days 13–14 on average and then receded in weight by 4%, apparently through loss of water. All healthy nestlings exceeded mean adult weight during development by up to 39%. Nestlings from broods of two were at first lighter on average than those from larger broods, but in the second half of the nestling period twins were significantly the heaviest. Grey Warblers were fed for 28–35 days after fledging and they survived well while dependent on parents. Fledglings dispersed up to 3 km or more at independence and only 5% per annum joined the breeding population. Of nests that received eggs, 42% produced at least one fledgling. On average each breeding adult raised 2·0 fledglings per season. Of 265 eggs in 73 nests 70% hatched and 38% produced fledglings. Of 185 nestlings 54% fledged. Probably the main cause of mortality of eggs and nestlings was predation by introduced rodents and mustelids. Grey Warblers raise two small broods slowly during a long breeding season, rather than investing in one large quickly-reared brood. In New Zealand's mild climate the warbler's food supply may not decline severely in winter, and the population of warblers may remain so close to the limit set by food that extra for breeding is hard to obtain. Thus the breeding strategy may be adapted to a restricted food supply.     

THE SOUTHWARD MIGRATION OF PALAEARCTIC BIRDS OVER NGULIA, KENYA

Since 1969 remarkable numbers of night migrants have been attracted during misty conditions in November and December to three 1 kW floodlights at a game viewing lodge on the northeastern side of the Ngulia ridge, a small range of hills in the Tsavo National Park (West), southeast Kenya. The main species involved have been Palaearctic passerines, principally the Marsh Warbler Acrocephalus palustris, the Whitethroat Sylvia communis and the Sprosser Luscinia luscinia. Data were collected at the Lodge in the late autumns of 1969–71, and in particular between November 1972 and early January 1973, when over 2500 Palaearctic passerines were caught and ringed. Large falls have depended on mist or rain during the latter part of the night, at any time during the month except around full moon. Highest numbers have occurred in late November and the first half of December. In 1973 falls continued into the second week of January. Grounded birds move on quickly, extremely few having been retrapped. During 1972–73, the species prominent in falls at the Lodge were abundant as transit migrants in Tsavo only from mid-December to early or mid January, at which time retrap rates were highest. The high weights and considerable fat deposits of many birds caught suggested they were grounded some distance north of their destination. Forty-two migrants analysed had a mean lipid content of 12·9% of their live weight; none was appreciably dehydrated. In 1972–73, highest weights were found at the beginning and end of the season. Individual species are discussed, and in several cases their African status reviewed. Several species were encountered at Ngulia in numbers far larger than those previously reported from elsewhere in Africa. In 1972–73, for instance, over 1000 Marsh Warblers were caught and many thousands of others seen, hundreds of River Warblers Locustella fluviatilis occurred, and White-throated Robins Irania gutturalis, Basra Reed Warblers Acrocephalus griseldis, Rufous Bush Chats Cercotrichas galactotes and Olive-tree Warblers Hippolais olivetorum were caught regularly. Most of the Basra Reed Warblers, Upcher's Warblers Hippolais languida and Olivaceous Warblers H. pallida, and many of the Whitethroats (apparently all of the eastern race icterops) handled during late December and early January were in fresh plumage, although these species are not known to moult north of the Sahara. They are presumed to have renewed their plumage in northeast Africa earlier in the autumn. In many Basra Reed Warblers and Whitethroats moult was only partly completed; in almost all such cases it was arrested. Itinerancy south of the Sahara is discussed. It seems clearly established that a regular southward migration, in the usually accepted sense of the word, occurs across Tsavo, of Palaearctic species which have already been in tropical Africa up to three months. Most species involved in this migration cross the equator on a remarkably narrow front, and are rarely recorded in Kenya west of Nairobi.     

Experimentally Constrained Virulence is Costly for Common Cuckoo Chicks

Chicks of some avian brood parasites show high virulence by eliminating all host progeny in the nest whereas others develop in the presence of host nestmates. Common cuckoo ( Cuculus canorus ) chicks are typically highly virulent parasites as they attempt to evict all host eggs and chicks soon after hatching. However, several features of nest design, including steep walls and/or cavity nests, may effectively prevent cuckoo hatchlings from evicting nestmates. A previous observational study showed low success of cuckoo chicks in evicting progeny of a cavity nester host, the redstart ( Phoenicurus phoenicurus ) but cuckoo chicks showed low survival both when reared alone or in mixed broods with host nestmates. Whether poor cuckoo performance was caused by eviction costs and/or by the effect of presence of host chicks per se remains unclear. We experimentally cancelled any potential eviction costs by removing host eggs immediately after the cuckoo hatched and creating mixed broods 5 days later when the eviction instinct of the cuckoo already ceased. Cuckoos that were forced to compete with host nestlings experienced lower provisioning rates, poorer growth, and lower fledging success than control lone cuckoos. Cuckoos in mixed broods that survived until fledging fledged later, and at lower masses, than those in the sole cuckoo group. Thus, the cuckoo gens specializing on redstarts is similar to other cuckoo gentes, whose chicks are more successful in evicting host nestmates, and it does not benefit from the presence of host brood. Cohabitation with host nestlings then should be viewed as a maladaptive by-product of host cavity nest design.     

Cryptic cuckoo eggs hide from competing cuckoos

Interspecific arms races between cuckoos and their hosts have produced remarkable examples of mimicry, with parasite eggs evolving to match host egg appearance and so evade removal by hosts. Certain bronze-cuckoo species, however, lay eggs that are cryptic rather than mimetic. These eggs are coated in a low luminance pigment that camouflages them within the dark interiors of hosts'' nests. We investigated whether cuckoo egg crypsis is likely to have arisen from the same coevolutionary processes known to favour egg mimicry. We added high and low luminance-painted eggs to the nests of large-billed gerygones (Gerygone magnirostris), a host of the little bronze-cuckoo (Chalcites minutillus). Gerygones rarely rejected either egg type, and did not reject natural cuckoo eggs. Cuckoos, by contrast, regularly removed an egg from clutches before laying their own and were five times more likely to remove a high luminance model than its low luminance counterpart. Given that we found one-third of all parasitized nests were exploited by multiple cuckoos, our results suggest that competition between cuckoos has been the key selective agent for egg crypsis. In such intraspecific arms races, crypsis may be favoured over mimicry because it can reduce the risk of egg removal to levels below chance.     

Fall migration and breeding origins of Canada Warblers moving through northern Colombia

Canada Warblers (Cardellina canadensis) are long‐distance Neotropical migrants, but little is known about their migratory behavior and ecology. We examined the fall migration of Canada Warblers at two sites, Darién and the Sierra Nevada de Santa Marta, in northern Colombia from 2011 to 2015 using constant‐effort mist‐netting. Our objectives were to determine: 1) breeding origins and connectivity patterns, 2) migratory pathways, 3) the phenology of migration, 4) possible differences in movements between ages and sexes, 5) their body condition when arriving in Colombia, and 6) evidence of stopover and refueling. Stable hydrogen isotopes (δ²H_f) in flight feathers were analyzed to estimate breeding origins of captured Canada Warblers in North America. The δ²H_f values revealed that most Canada Warblers captured in the Darién likely originated from the central and northeastern regions of their breeding range. The capture of all but one of 162 Canada Warblers in the Darién also indicates a migration route through Central American rather than across the Caribbean Sea. Most captured birds were hatch‐year birds (91% vs. 9% after hatch‐year birds), and we captured more females (67%) than males (33%). Canada Warblers migrated through the Darién from 20 September to early November, with most arriving in mid‐October. Most (89%) individuals arrived with low fuel reserves. These results combined with estimated flight ranges revealed that 46% of the individuals captured in the Darién likely needed to refuel to continue migrating, whereas 31% could continue 50 to 200 km beyond our capture site. However, no individuals were recaptured so stopover duration could not be determined. Canada Warblers may adopt a strategy of 1‐d stopovers and short flights or, alternatively, the Darién may represent low‐quality habitat and birds quickly left our study site in search of suitable habitat. Further study is needed to determine the possible importance of other (montane) habitats for Canada Warblers in the Darién region to prioritize conservation actions.     

When should Common Cuckoos Cuculus canorus lay their eggs in host nests?

Capsule Brood parasitic Common Cuckoo Cuculus canorus chicks hatch earlier than the nestlings of their Great Reed Warbler Acrocephalus arundinaceus hosts, but hatching priority is less consistent when Cuckoo eggs are laid after the onset of host incubation.

Aim To reveal by field observations what the optimal stage is for Cuckoos to lay their eggs in relation to the host laying cycle to ensure prior hatching of the parasitic chicks.

Methods We monitored the hatching of Cuckoo chicks in relation to the hosts’ laying stage at which the Cuckoo eggs appeared and also monitored host incubation behaviour.

Results Great Reed Warblers incubated more on day 5 after the host's onset of laying relative to day 3. All Cuckoo eggs hatched earlier than hosts when they were laid prior to the onset of host incubation (day 4). Cuckoo eggs also maintained hatching priority in about 2/3 of the nests when laid on days 5–6.

Conclusions Most Cuckoo eggs are laid prior to the onset of host incubation and this, together with other adaptive mechanisms, ensures the prior hatching of Cuckoo eggs. Cuckoo eggs laid after the onset of incubation lose the advantage of prior hatching in approximately 30% of nests.     

Behaviour of female common cuckoos, Cuculus canorus, in the vicinity of host nests before and during egg laying: a radiotelemetry study

We radiotracked 13 common cuckoo females in the southeastern part of the Czech Republic. Seven females laid eggs in the nests of reed warblers, Acrocephalus scirpaceus, sedge warblers, A. schoenobaenus, and marsh warblers, A. palustris. We observed 53 nest visits, of which 26 involved egg laying. Cuckoos spent significantly more time within 50 m of the host nest on the laying day than on the 5 prelaying days. The vantage point used when parasitizing or visiting a nest was on average four times further from the nest than the closest possible vantage point, but there was a positive correlation between these two distances. Cuckoos spent on average 20 min observing host nests from their vantage points before they visited a nest. Comparison of cuckoos' visits to host nests with and without egg laying revealed no significant differences in the duration of visits or in other measures of behaviour. There was significant variation in behaviour between cuckoos, particularly in the time of day when eggs were laid in host nests. This variation can be attributed to the strong, but not absolute, host and habitat specificity of individuals. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Predicting the responses of native birds to transoceanic invasions by avian brood parasites

Three species of brood parasites are increasingly being recorded as transoceanic vagrants in the Northern Hemisphere, including two Cuculus cuckoos from Asia to North America and a Molothrus cowbird from North America to Eurasia. Vagrancy patterns suggest that their establishment on new continents is feasible, possibly as a consequence of recent range increases in response to a warming climate. The impacts of invasive brood parasites are predicted to differ between continents because many host species of cowbirds in North America lack egg rejection defenses against native and presumably also against invasive parasites, whereas many hosts of Eurasian cuckoos frequently reject non‐mimetic, and even some mimetic, parasitic eggs from their nests. During the 2014 breeding season, we tested the responses of native egg‐rejecter songbirds to model eggs matching in size and color the eggs of two potentially invasive brood parasites. American Robins (Turdus migratorius) are among the few rejecters of the eggs of Brown‐headed Cowbirds (M. ater), sympatric brood parasites. In our experiments, robins rejected one type of model eggs of a Common Cuckoo (C. canorus) host‐race, but accepted model eggs of a second cuckoo host‐race as well as robin‐mimetic control eggs. Common Redstarts (Phoenicurus phoenicurus), frequent hosts of Common Cuckoos in Eurasia, rejected ～50% of model Brown‐headed Cowbird eggs and accepted most redstart‐mimetic control eggs. Our results suggest that even though some hosts have evolved egg‐rejection defenses against native brood parasites, the invasion of brood parasites into new continents may negatively impact both naïve accepter and coevolved rejecter songbirds in the Northern Hemisphere.     

Responses of potential hosts of Asian cuckoos to experimental parasitism

In the arms race between avian brood parasites and their hosts, several adaptations and counter‐adaptations have evolved. The most prominent host defence is rejection of parasitic eggs. We experimentally parasitized nests of 10 potential host species breeding in sympatry with four different cuckoo species in an area in Bangladesh using differently coloured model eggs to test host responses. In four species we introduced both mimetic and non‐mimetic eggs. Black Drongos Dicrurus macrocercus, hosts of the Indian Cuckoo Cuculus micropterus, rejected all model eggs. Common Mynas Acridotheres tristis and Jungle Babblers Turdoides striata accepted all eggs regardless of mimicry. These two species are parasitized by Asian Koels Eudynamys scolopaceus, Common Hawk‐cuckoo Hierococcyx varius and, in the case of Jungle Babblers, Jacobin Cuckoos Clamator jacobinus. Pied Mynas Gracupica contra, with no records of parasitism in our study area, also accepted all eggs regardless of mimicry. In the six remaining species, all of which lay spotted eggs, we introduced only non‐mimetic eggs. Black‐hooded Orioles Oriolus xanthornus rejected all model eggs, even though we have found no records of natural parasitism. Long‐tailed Shrikes Lanius schach and House Crows Corvus splendens, hosts of Asian Koels, rejected 75 and 9.1% of model eggs, respectively. Large‐billed Crows Corvus macrorhynchos, apparently not used as hosts in our study area, accepted all blue but rejected all brown model eggs. Oriental Magpie‐Robins Copsychus saularis and Red‐vented Bulbuls Pycnonotus cafer accepted all non‐mimetic model eggs. In Black Drongos, Long‐tailed Shrikes and Black‐hooded Orioles, all model eggs were ejected within 24 h of introduction. The results show considerable variation in egg rejection rates among various species, providing baseline data for further investigation of co‐evolutionary interactions between brood parasites and hosts in this region.     

Dawn singing of the Brownish‐flanked Bush Warbler influences dawn chorusing in a bird community

Despite numerous studies on the function of the avian dawn chorus, few studies have examined whether dawn singing may influence the singing of other species. Here, we built on our previous study which found male Brownish‐flanked Bush Warblers (Horornis fortipes) increase their dawn singing intensity after conspecific playback on the previous day. We reanalyzed those recordings to quantify the start of dawn singing in other nine sympatric songbird species. Ranking‐scaling analyses identified a distinctive sequential pattern of dawn singing among these bird species between the first and the second dawn chorus, and meta‐analysis showed a significant trend to singing earlier in the bird community accompanied by the increase in dawn singing intensity in Brownish‐flanked Bush Warbler. Species with songs most similar to that of the Brownish‐flanked Bush Warbler and species that were phylogenetically distantly related to the Brownish‐flanked Bush Warbler showed a greater shift in the onset of dawn singing. Our study is one of the few studies showing how bird song influences heterospecific singing, and this may influence the temporal organization of song activity in the community, and result in synchronization in singing activities among different species, such as singing in dawn and dusk chorus.