Common cuckoo

The common cuckoo's diet consists of insects, with hairy caterpillars, which are distasteful to many birds, being a specialty of preference.[14] It also occasionally eats eggs and chicks.[15]

More than 100 host species have been recorded: meadow pipit, dunnock and Eurasian reed warbler are the most common hosts in northern Europe; garden warbler, meadow pipit, pied wagtail and European robin in central Europe; brambling and common redstart in Finland; and great reed warbler in Hungary.[4]

Female common cuckoos are divided into gentes – groups of females favouring a particular host species' nest and laying eggs that match those of that species in color and pattern. Evidence from mitochondrial DNA analyses suggest that each gente may have multiple independent origins due to parasitism of specific hosts by different ancestors.[17] One hypothesis for the inheritance of egg appearance mimicry is that this trait is inherited from the female only, suggesting that it is carried on the sex-determining W chromosome (females are WZ, males ZZ). A genetic analysis of gentes supports this proposal by finding significant differentiation in mitochondrial DNA, but not in microsatellite DNA.[17] A second proposal for the inheritance of this trait is that the genes controlling egg characteristics are carried on autosomes rather than just the W chromosome. Another genetic analysis of sympatric gentes supports this second proposal by finding significant genetic differentiation in both microsatellite DNA and mitochondrial DNA.[18] Considering the tendency for common cuckoo males to mate with multiple females and produce offspring raised by more than one host species, it appears as though males do not contribute to the maintenance of common cuckoo gentes. However, it was found that only nine percent of offspring were raised outside of their father's presumed host species.[18] Therefore, both males and females may contribute to the maintenance of common cuckoo egg mimicry polymorphism.[17][18] It is notable that most non-parasitic cuckoo species lay white eggs, like most non-passerines other than ground-nesters.

Cuckoo eggs mimicking smaller eggs, in this case of reed warbler

Cuculus canorus canorus in a nest Acrocephalus arundinaceus - MHNT

Cuculus canorus bangsi in a nest Phoenicurus moussieri - MHNT

As the common cuckoo evolves to lay eggs that better imitate the host's eggs, the host species adapts and is more able to distinguish the cuckoo egg. A study of 248 common cuckoo and host eggs demonstrated that female cuckoos that parasitised common redstart nests laid eggs that matched better than those that targeted dunnocks. Spectroscopy was used to model how the host species saw the cuckoo eggs. Cuckoos that target dunnock nests lay white, brown-speckled eggs, in contrast to the dunnock's own blue eggs. The theory suggests that common redstarts have been parasitised by common cuckoos for longer, and so have evolved to be better than the dunnocks at noticing the cuckoo eggs. The cuckoo, over time, has needed to evolve more accurate mimicking eggs to successfully parasitise the redstart. In contrast, cuckoos do not seem to have experienced evolutionary pressure to develop eggs which closely mimic the dunnock's, as dunnocks do not seem to be able to distinguish between the two species' eggs, despite the significant colour differences. The dunnock's inability to distinguish the eggs suggests that they have not been parasitised for very long, and have not yet evolved defences against it, unlike the redstart.[19]

Studies performed on great reed warbler nests in central Hungary, showed an "unusually high" frequency of common cuckoo parasitism, with 64% of the nests parasitised. Of the nests targeted by cuckoos, 64% contained one cuckoo egg, 23% had two, 10% had three and 3% had four common cuckoo eggs. In total, 58% of the common cuckoo eggs were laid in nests that were multiply parasitised. When laying eggs in nests already parasitised, the female cuckoos removed one egg at random, showing no discrimination between the great reed warbler eggs and those of other cuckoos.[20]

It was found that nests close to cuckoo perches were most vulnerable: multiple parasitised nests were closest to the vantage points, and unparasitised nests were farthest away. Nearly all the nests "in close vicinity" to the vantage points were parasitised. More visible nests were more likely to be selected by the common cuckoos. Female cuckoos use their vantage points to watch for potential hosts and find it easier to locate the more visible nests while they are egg-laying,[21] however, novel studies highlight that host alarm calls might also play an important role during nest searching.[22] In addition, cuckoos tend to lay the eggs on the host clutch initiation day or one day before.[23]

The great reed warblers' responses to the common cuckoo eggs varied: 66% accepted the egg(s); 12% ejected them; 20% abandoned the nests entirely; 2% buried the eggs. 28% of the cuckoo eggs were described as "almost perfect" in their mimesis of the host eggs, and the warblers rejected "poorly mimetic" cuckoo eggs more often. The degree of mimicry made it difficult for both the great reed warblers and the observers to tell the eggs apart.[20]

The egg measures 22 by 16 millimetres (0.87 in × 0.63 in) and weighs 3.2 grams (0.11 oz), of which 7% is shell.[2] Research has shown that the female common cuckoo is able to keep its egg inside its body for an extra 24 hours before laying it in a host's nest. This means the cuckoo chick can hatch before the host's chicks do, and it can eject the unhatched eggs from the nest. Scientists incubated common cuckoo eggs for 24 hours at the bird's body temperature of 40 °C (104 °F), and examined the embryos, which were found "much more advanced" than those of other species studied. The idea of 'internal incubation' was first put forward in 1802 and 18th- and 19th-century egg collectors had reported finding that cuckoo embryos were more advanced than those of the host species.[24]

A study using digital photography and spectrometry along with an automatic analytical approach to analyse cuckoo eggs and predict the identity of bird females based on their egg appearance showed that individual cuckoo females lay eggs with a relatively constant appearance, and that eggs laid by more genetically distant females differ more in colour.[25]

Complete list of common cuckoo's nest-host by Aleksander D. Numerov (2003); names of birds in whose nests cuckoo's eggs and chicks were found more than 10 times (in bold):[26]

A chick of the common cuckoo in the nest of a tree pipit

The naked, altricial chick hatches after 11–13 days.[2] It methodically evicts all host progeny from host nests. It is a much larger bird than its hosts, and needs to monopolize the food supplied by the parents. The chick will roll the other eggs out of the nest by pushing them with its back over the edge. If the host's eggs hatch before the cuckoo's, the cuckoo chick will push the other chicks out of the nest in a similar way. At 14 days old, the common cuckoo chick is about three times the size of an adult Eurasian reed warbler.

The necessity of eviction behavior is unclear. One hypothesis is that competing with host chicks leads to decreased cuckoo chick weight, which is selective pressure for eviction behavior. An analysis of the amount of food provided to common cuckoo chicks by host parents in the presence and absence of host siblings showed that when competing against host siblings, cuckoo chicks did not receive enough food, showing an inability to compete.[33] Selection pressure for eviction behavior may come from cuckoo chicks lacking the correct visual begging signals, hosts distributing food to all nestlings equally, or host recognition of the parasite.[33][34] Another hypothesis is that decreased cuckoo chick weight is not selective pressure for eviction behavior. An analysis of resources provided to cuckoo chicks in the presence and absence of host siblings also showed that the weights of cuckoos raised with host chicks were much smaller upon fledging than cuckoos raised alone, but within 12 days cuckoos raised with siblings grew faster than cuckoos raised alone and made up for developmental differences, showing a flexibility that would not necessarily select for eviction behavior.[35]

Species whose broods are parasitised by the common cuckoo have evolved to discriminate against cuckoo eggs but not chicks.[36] Experiments have shown that common cuckoo chicks persuade their host parents to feed them by making a rapid begging call that sounds "remarkably like a whole brood of host chicks." The researchers suggested that "the cuckoo needs vocal trickery to stimulate adequate care to compensate for the fact that it presents a visual stimulus of just one gape."[34] However, a cuckoo chick needs the amount of food of a whole brood of host nestlings, and it struggles to elicit that much from the host parents with only the vocal stimulus. This may reflect a tradeoff—the cuckoo chick benefits from eviction by receiving all the food provided, but faces a cost in being the only one influencing feeding rate. For this reason, cuckoo chicks exploit host parental care by remaining with the host parent longer than host chicks do, both before and after fledging.[34]

Common cuckoo chicks fledge about 17–21 days after hatching,[2] compared to 12–13 days for Eurasian reed warblers.[37] If the hen cuckoo is out-of-phase with a clutch of Eurasian reed warbler eggs, she will eat them all so that the hosts are forced to start another brood.

The common cuckoo's behaviour was firstly observed and described by Aristotle and the combination of behaviour and anatomical adaptation by Edward Jenner, who was elected as Fellow of the Royal Society in 1788 for this work rather than for his development of the smallpox vaccine. It was first documented on film in 1922 by Edgar Chance and Oliver G Pike, in their film The Cuckoo's Secret.[38] A study in Japan found that young common cuckoos probably acquire species-specific feather lice from body-to-body contact with other cuckoos between the time of leaving the nest and returning to the breeding area in spring. A total of 21 nestlings were examined shortly before they left their hosts' nests and none carried feather lice. However, young birds returning to Japan for the first time were found just as likely as older individuals to be lousy.[39]