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To maximize their offspring success common cuckoo ( Cuculus canorus ) females should lay their eggs into host nests before incubation has begun. This ensures that the parasite chick hatches before all host chicks and can evict its foster siblings to monopolize host parental care. Many studies have demonstrated that most cuckoo eggs are indeed laid before the onset of host incubation. But cues used by female cuckoos to choose the right nest at the right time remain unclear. Here, we combine field observations with a field experiment to test whether female cuckoos use the number of eggs in the nests of their Oriental reed warbler ( Acrocephalus orientalis ) hosts to direct their choice. Over 8 years of field observations and 5 years of experiments, cuckoo females placed the majority of their eggs in nests with fewer than three host eggs, i.e. early in the laying sequence. For natural nests, the cuckoos may use information gleaned from the activity and behaviour of the host parents to make their choice. In our sets of experimental nests containing different numbers of model eggs, the vast majority of parasitism events occurred in nests containing a single egg. To our knowledge, this is the first field experiment showing that cuckoos choose host nests for parasitism based on the number of host eggs they contain. It appears that cuckoo females use the egg number to estimate the appropriate host nest stage for timely parasitism.

a: Cuckoo flew down to target nest; b: Host guarded nest; c: Cuckoo perched on target nest; d: Cuckoo hid its head and started egg pecking/biting; e: Cuckoo hunched its body, pecked an egg, closed its tail, and opened its eyes; f: Cuckoo finished egg laying, sometimes removing an egg; g: Cuckoo flew away.

Nest defense is an effective strategy of hosts against parasites. Typically, hosts will aggressively attack brood parasites that approach or visit their nests, which can prevent the parasites from laying eggs or may even lead to the death of the parasites. Few previous studies have specifically reported such fatal cases involving brood parasites and have attributed the cause of death to either drowning or hypothermia after falling into the water following an attack from hosts. In this study, we recorded the process of multiple host individuals of the Oriental reed warbler (Acrocephalus orientalis) mobbing and attacking a female common cuckoo (Cuculus canorus) in the field. We discovered that the immediate cause of the cuckoo's death was the fatal physical damage resulting from the aggressive defense from the hosts, suggesting that frantic pecking and scratching by the hosts is the most proximate cause of mortality among egg‐laying female cuckoos. This finding enhances our essential understanding of the effectiveness of host attacks.

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