Obligate brood parasites rely solely on other species, the hosts, to incubate their eggs and raise their offspring, which often reduces the host’s reproductive output. This reproductive cost has led to the evolution of anti-parasite adaptations among hosts, which in turn, has led to better trickery by parasites, a process termed an evolutionary arms race. The objective of this thesis was to investigate host-parasite coevolutionary arms races to address questions of host-use diversity. Host diversity varies dramatically among brood-parasitic species, but reasons for variations in host-use among brood parasites are not well understood. In Chapter 2, I address questions on host diversity specifically, whereas I address questions about coevolutionary interaction between hosts and parasites in Chapters 3, 4 and 5 using two host-parasite systems, one in New Zealand and one in North America. Chapter 2 investigates if host diversity is constrained by aggressive nest defence behaviour. I compared the nest defence behaviour of the exclusive host of the shining cuckoo Chrysococcyx lucidus lucidus on the main islands of New Zealand, the grey warbler Gerygone igata, to two other potentially suitable hosts that are not currently parasitised, the fantail Rhipidura fuliginosa and the silvereye Zosterops lateralis. The results suggest that grey warblers are as aggressive as fantails and silvereyes towards shining cuckoos at the nest and thus, host specialisation in shining cuckoos in New Zealand, at least, does not appear to be the result of nest-defence constraints imposed by potential but unused host species. Chapter 3 investigates if red-winged blackbirds Agelaius phoeniceus, a species that typically accepts the eggs of parasites, recognises, as indicated by changes in incubation behaviour, when they have been parasitised by brown-headed cowbirds Molothrus ater. Recognition without rejection suggests that rejection may be context-dependent but the results suggest that red-winged blackbirds do not recognise when their nests have been parasitised by brown-headed cowbirds, at least at the egg stage. This study was the first to investigate if hosts that almost invariably accept the eggs of parasites recognise when they have been parasitised. Chapter 4 investigated the possibility of coevolutionary arms races occurring through olfactory channels in contrast to earlier work that focussed only on visual and auditory cues. Recent research has revealed that olfactory abilities in birds are more common than previously thought. Uropygial gland secretions are posited to be a key source of avian body odour and its composition has been found to vary among species and individuals as well as between the sexes. I compared gas-chromatography (GC-FID) traces of shining cuckoo preen wax to the GC-FID traces of the grey warbler, the only host of the shining cuckoo in mainland New Zealand, as well as the preen wax of seven other species for evidence of mimicry. Preliminary results suggest there is evidence for mimicry and the potential for odour-based nestling discrimination in grey warblers. Further tests recording the response of grey warblers to odour-manipulated nestlings are necessary. Finally, in Chapter 5, I investigated the response of the song thrush Turdus philomelos, a species that rejects the eggs of the common cuckoo Cuculus canorus and conspecifics at intermediate and low frequencies, respectively, to nest-odour manipulations using the preen wax of conspecifics and heterospecifics. The results suggest song thrush do not use odour to assess the risk of parasitism at least as indicated in terms of changes in incubation behaviour. Investigations of the role of olfaction in avian brood parasite systems can provide a better understanding of brood-parasite coevolution. Only by considering all channels of communication can we be sure to completely understand the coevolutionary dynamics between brood parasites and their hosts.

Obligate brood parasites lay their eggs in nests of other species, relying on these host parents to care for their offspring. This phenomenon has been a curiosity amongst researchers since its first description and has become a model study system for testing such ideas as coevolution and species recognition. This thesis examines a few of the many questions that arise from this breeding system. The New Zealand Grey Warbler (Gerygone igata) and its brood parasite, the Shining Cuckoo (Chrysococcyx lucidus) are used as the main study species, although research on the eviction behaviour of Common Cuckoos (Cuculus canorus) has also been conducted. First, the current state of knowledge and recent discoveries regarding nestling rejection abilities of hosts is reviewed in chapter one. Second, a comparative study of New Zealand passerine begging calls has been conducted to test for begging call similarity between a brood parasite and its host, as well as developing a new technique for detecting the mode of coevolution that may be occurring in the parasite – host relationship. Parent-offspring communication in Grey Warblers is also examined to test for both parental and nestlings Parents use both alarm calls to warn offspring of potential danger, and also parental feeding calls to elicit a begging response from nestlings. By contrast, nestlings are able to signal both age and short term levels of need to parents through the acoustic structure of the begging call. The evolutionary costs and benefits of egg eviction behaviour in the Common Cuckoo are also tested. An experimental approach showed that egg eviction had a growth cost, but this cost was temporary and restricted to during and immediately after the egg eviction phase. A pattern of compensatory growth was observed after the eviction period, so that during the later nestling stages there was no difference in mass, and no difference in fledging age. Finally, variation in the Grey Warbler breeding biology and Shining Cuckoo parasitism rates are examined through both time and across latitudes. This research has shown a counterintuitive pattern of breeding phenology across latitudes. These patterns have implications for Shining Cuckoos both in terms of timing of available nests and host selection. Keywords: Begging call, breeding phenology, brood parasitism, coevolution, Common Cuckoo, eviction, Grey Warbler, parent-offspring communication, Shining Cuckoo.