Brood parasitic birds lay eggs amongst the clutches of other species, which then assume all costs of parental care on their behalf. This thesis addresses several puzzles of avian brood parasitism, using field studies and theoretical modelling of the generalist parasite, the shiny cowbird (Molothrus bonariensis) and select hosts in Argentina. Key findings and conclusions were: • High parasitism intensity in a host population can result in a cost to hosts of removing parasite eggs from their clutches, and so help to maintain host’s acceptance of parasite eggs in evolutionary equilibrium. The cost is to host egg survival: hosts that remove parasite eggs from the clutch increase the risk that their eggs are destroyed by subsequent parasites that visit the nest. • The principal benefit of mobbing as a front-line defence of hosts may be to reduce egg loss due to parasite attack, rather than prevent parasitism itself. • Differences in the acoustic structure of begging calls between parasites and their host’s young can be to the parasite’s advantage. Parents provisioned unparasitized broods more during broadcast at the nest of shiny cowbird calls than calls of their own species’ chicks, in both a common host and a non-host. The long tremulous quality of a cowbird’s call functions analogously to a rapid call rate, thereby exploiting a common provisioning rule of avian parents. • A trade-off for maximum growth in parasite nestlings will variously favour or not favour the evolution of nestmate-killing behaviour, depending on a parasite’s abilities, relative to host young, to solicit and attain provisions from host parents. Generalist parasites can encounter both sides of the trade-off in different hosts. Meanwhile, indirect fitness costs are unlikely to constrain the evolution of nestmate-killing in shiny cowbirds, as they rarely, if ever, share the nest with siblings.

The proximate causes triggering nest abandonment are unclear for most species, including the Yellow Warbler (Dendroica petechia), which abandons nests parasitized by cowbirds (via burial or desertion). Cowbird parasitism and rejection of parasitism are costly to some hosts; therefore cues affecting their responses have important evolutionary implications. Manipulative experiments showed that experimentally adding a cowbird egg elicited similar rejection frequencies (2008: 31.8%; 2009: 26.1%) as naturally laid eggs (2008: 27.1%; 2009: 20.0%). In 2008, interaction with an egg-removing model increased the probability of abandonment and the most aggressive individuals were more likely to bury the model cowbird egg. In 2009, eggs added to nests before sunrise were rejected at a frequency (29.7%) similar to eggs added to nests after sunrise (22.9%). Warblers returning to nests after egg addition peered significantly longer at their clutch than at control nests, shuffled their bodies more frequently when on the eggs and spent more time probing eggs with their bill once settled on their parasitized clutch. Furthermore, although non-mimetic blue eggs were not abandoned significantly more frequently than cowbird eggs (blue 31.1% versus cowbird 21.4%), only blue eggs were ejected from some nests. Thus, warblers use both tactile and visual cues to detect the presence of a parasitic egg in their nest. Eggs added to nests were not rejected at a lower frequency than naturally parasitized nests, as was recorded in a previous study. It is difficult to know whether this increase in abandonment of experimental eggs is due to phenotypic plasticity, genetic changes, or other factors. Egg recognition abilities may have changed because I have shown that the warblers’ behaviour changes before versus after egg addition, whereas no changes were recorded in an earlier study. Finally, not all individuals that buried eggs for the first time in 2009 (21.4%) buried again after being re-parasitized (5.3%), when less time remained in the breeding season relative to the first parasitism event. This suggests that egg rejection and host responsiveness in warblers, and likely other avian hosts that use abandonment as a form of rejection, is affected by environmental cues which may act as genetic expressers.

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.

The proximate causes triggering nest abandonment are unclear for most species, including the Yellow Warbler (Dendroica petechia), which abandons nests parasitized by cowbirds (via burial or desertion). Cowbird parasitism and rejection of parasitism are costly to some hosts; therefore cues affecting their responses have important evolutionary implications. Manipulative experiments showed that experimentally adding a cowbird egg elicited similar rejection frequencies (2008: 31.8%; 2009: 26.1%) as naturally laid eggs (2008: 27.1%; 2009: 20.0%). In 2008, interaction with an egg-removing model increased the probability of abandonment and the most aggressive individuals were more likely to bury the model cowbird egg. In 2009, eggs added to nests before sunrise were rejected at a frequency (29.7%) similar to eggs added to nests after sunrise (22.9%). Warblers returning to nests after egg addition peered significantly longer at their clutch than at control nests, shuffled their bodies more frequently when on the eggs and spent more time probing eggs with their bill once settled on their parasitized clutch. Furthermore, although non-mimetic blue eggs were not abandoned significantly more frequently than cowbird eggs (blue 31.1% versus cowbird 21.4%), only blue eggs were ejected from some nests. Thus, warblers use both tactile and visual cues to detect the presence of a parasitic egg in their nest. Eggs added to nests were not rejected at a lower frequency than naturally parasitized nests, as was recorded in a previous study. It is difficult to know whether this increase in abandonment of experimental eggs is due to phenotypic plasticity, genetic changes, or other factors. Egg recognition abilities may have changed because I have shown that the warblers’ behaviour changes before versus after egg addition, whereas no changes were recorded in an earlier study. Finally, not all individuals that buried eggs for the first time in 2009 (21.4%) buried again after being re-parasitized (5.3%), when less time remained in the breeding season relative to the first parasitism event. This suggests that egg rejection and host responsiveness in warblers, and likely other avian hosts that use abandonment as a form of rejection, is affected by environmental cues which may act as genetic expressers.

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.

Conspecific brood parasitism (CBP) is a situation, when female (a parasite) lays egg or eggs in a nest of other conspecific female (a host). Then she leaves the nest and doesn't provide any subsequent parental care, such incubation and care for young. To distinguish between parasitic and host egg or young could be difficult, because there are not obvious morphological differences. CBP was documented in more than 234 species and its frequency is different between species, but also between population of the same species. The main aim of my master thesis was to describe the degree of parasitism in two population of Common Pochard (Aythya ferina). The field work was carried out from 2004-2006 in Třeboň Biosphere Reserve and form 2008-2010 in Poodří Landscape Protected Area. We found that the degree of CBP is relatively high. In Třeboň was found 72 % (41) of parasitized nests and 32 % (165) of parasitic eggs and in Poodří 93 % (39) of parasitized nests and 37 % (153) of parasitic eggs. The mean number of eggs per clutch was around 9 and mean number of host eggs per clutch was 6 in Třeboň. The mean number of eggs per clutch in Poodří was about 10, and the mean number of host eggs was also about 6. The mean number of parasitic eggs per clutch was 3 resp. 4. It was also found, that parasitized nests are...

This study is aimed at analysis of long-term and short-term changes in numbers of Red-crested Pochard (Netta rufina) in Třeboň region. This species is increasing in long-term period and it is also spreading into new localities in the Czech Republic. There were recorded two waves of arrival of Red-crested Pochard arrive, i.e. in April and in May. The total numbers were decreasing at the end of breeding season (from the beginning of July) in study area. The water surface area and area of littoral vegetation were found to be the most important factors affecting preference of individual ponds in pre-breeding season. Furthermore, water transparency seems to be important factor affecting numbers of broods, ducklings and adults in post-breeding season. Low numbers of broods per 1 female were recorded in years with high numbers of Red- crested Pochard at the start of breeding season. However, the frequency of brood parasitism was higher in these years. Key words: Netta rufina, Red-crested Pochard, population dynamics, ducks, breeding, numbers, brood parasitism, south Bohemia

Avian brood parasitism is an ideal system for the study of coevolution. Brood parasites and their hosts have developed interesting adaptations during co-evolution allowing them to maximize their fitness. The evolution of these adaptations has a character of an "arms race" where the evolution of one trait in the host is tied with the evolution of another trait in the parasite. In my doctoral thesis, I deal with two of these adaptations: recognition of parasitic eggs by hosts and mimicry of eggs in parasites. Since both these adaptations are influenced by birds' visual system, in all my studies I used an objective method to measure the colour and the modelling of avian visual system that is quite different from the human visual system. For instance, humans in contrast to birds cannot perceive ultraviolet (UV) light. However, this part of spectrum influences behaviour of birds substantially (e.g. courtship or foraging). We found that the hosts of brood parasites can use UV light when recognizing parasitic eggs. However, it seems that this part of spectrum is not the main cue in egg recognition (manuscript 1). Ambient light has also an important impact on colour perception. We determined whether the light conditions in nests influence host responses to alien eggs. The Red Bishop (Euplectes orix) was an ideal...

Abstract Animals live in a heterogeneous world where threats and abundance and quality of resources vary across space and time. Heterogeneity induces uncertainty in decisions that animals must make, e.g., where to breed. Adaptive decisions may be facilitated by personally collecting information on the quality of the environment and by observing the behaviour and success of other individuals. Such social information use is common in nature. I investigate information use in relation to ecological threats (brood parasites, nest predators) and long-term information use in breeding site choice in the wild. Moreover, I examine the genetic basis of social cue use in breeding site choice. I demonstrated experimentally that open-nesting hosts of a brood parasite, the common cuckoo (Cuculus canorus), can cue on cuckoo vocalizations to estimate cuckoo abundance and avoid breeding sites with high perceived parasitism risk. Another experiment showed that pied flycatchers (Ficedula hypoleuca) derive predation risk information from the fates of heterospecific nests, can associate the information with a nest site characteristic and generalize the association to own nest site choice. However, apparently only young females that made their choice quickly used the information in nest site choice. Pied flycatchers were further observed to collect habitat quality information based on the old nest contents during the post-breeding period. Use of the information in breeding site choice in the following spring varied between sex and age groups as well as geographically. Some birds integrated the post-breeding period information with the information available during settlement suggesting sequential social information use. Finally, quantitative genetic analyses revealed low additive genetic variances and genetic heritabilities of social cue use in breeding site choice in a collared flycatcher (Ficedula albicollis) population. These results demonstrate new aspects of informed habitat choice in wild animals which have important implications for species coexistence and community ecology, parasite-host coevolution, between-species niche dynamics and evolution. Between-individual variation in information use is highlighted throughout the thesis and warrants further research. The evolutionary potential of information use appears low, but more studies in other populations and species are needed. / Tiivistelmä Eläimet elävät ympäristössä, jossa resurssit ja uhat vaihtelevat ajallisesti ja alueellisesti. Tämä vaihtelu aiheuttaa epävarmuutta eläinten päätöksentekoon, kuten pesimäpaikan valintaan. Hyödyllisten päätösten tekoa voi edesauttaa keräämällä tietoa ympäristön laadusta itsenäisesti tai seuraamalla muiden yksilöiden käytöstä ja menestystä. Tällainen sosiaalisen informaation käyttö on yleistä eläinkunnassa. Tutkin informaation käyttöä ekologisten uhkien (pesäloiset, -pedot) suhteen ja pitkäaikaista informaation käyttöä pesimäpaikan valinnassa luonnonpopulaatioissa. Lisäksi selvitän pesimäpaikan valintaan liittyvän informaation käyton geneettistä periytyvyyttä. Selvitin kokeellisesti, että pesäloisen, käen (Cuculus canorus), isäntälajit voivat käyttää käkien ääntelyä vihjeenä alueellisesta loisintauhasta ja siten välttää korkean uhan alueita pesimäpaikan valinnassa. Toisessa kokeessa havaittiin kirjosieppojen (Ficedula hypoleuca) keräävän tietoa pesäpetouhasta toisen lajin pesätuhojen kautta, kykenevän yhdistämään tiedon erilliseen pesäpaikan ominaisuuteen ja käyttämään tätä assosiaatiota omassa pesäpaikan valinnassa. Kuitenkin vain nuoret naaraat, jotka tekivät valintansa nopeasti, käyttivät kyseistä informaatiota valinnassaan. Lisäksi havaitsin kirjosieppojen keräävän tietoa ympäristön laadusta pesinnän jälkeen vanhojen pesäsisältöjen avulla. Kyseisen tiedon käyttö pesimäpaikan valinnassa seuraavana keväänä vaihteli lintujen sukupuolen ja iän suhteen, kuin myös alueellisesti. Osa linnuista yhdisti pesimäpaikan valinnassaan aikaisempaa, pesinnän jälkeen kerättyä tietoa ja keväällä saatavilla olevaa sosiaalista informaatiota. Geneettinen analyysi viittasi pesimäpaikan valintaan liittyvän informaation käytön alhaiseen additiivisen geneettisen varianssin määrään ja siten alhaiseen geneettiseen periytyvyyteen sepelsiepolla (Ficedula albicollis). Väitöskirjani tulokset kuvaavat uudenlaisia informaation käytön muotoja eläinten pesimäpaikan valinnassa. Havainnot auttavat ymmärtämään pesälois-isäntä rinnakkaisevoluutiota, lajien välisiä vuorovaikutuksia, lajiyhteisöjen toimintaa ja evoluutiota. Yksilöiden välinen vaihtelu informaation käytössä näyttää olevan yleistä, ja lisätutkimuksen tarpeessa. Informaation käytön evolutiivinen potentiaali näyttää rajalliselta, mutta lisätutkimukset eri populaatioilla ja lajeilla ovat tarpeen.

Doctor of Philosophy / Department of Biology / Brett K. Sandercock / The reproductive strategies of avian brood parasites and the behavioral responses of their hosts have served as a model of co-evolution in nature. Host adaptations to reduce the costs of parasitism are countered with novel parasite behaviors that increase the success of the parasite and thereby decrease host productivity. Not all host species possess anti-parasite defense behaviors, and parasitism by Brown-headed Cowbirds (Molothrus ater) may cause population declines in some species. Bell’s Vireo (Vireo bellii) is a small-bodied cowbird host that fails to fledge young if successfully parasitized. Although vireos desert naturally parasitized nests, the cues that cause desertion have not been identified. Understanding how parasitism affects vireo productivity is important because cowbird removal is an integral component of the recovery efforts for the endangered Least Bell’s Vireo (V. b. pusillus) in California. However, it is generally unknown how cowbird removal affects vireo productivity. To address these issues, I monitored the productivity of vireos nesting in Kansas at the Konza Prairie Biological Station, conducted a clutch manipulation experiment, and experimentally removed cowbirds. In addition, I used stable isotope analysis to determine if recently fledged cowbird young could be assigned to habitats or host species. I found that vireos did not desert nests due to the presence of a cowbird egg; rather egg removal by cowbirds caused desertion, which is a generalized response in many taxa of birds. Cowbird removals decreased parasitism of vireo nests by approximately 36% and led to a 2-fold increase in vireo productivity per pair. Cowbird productivity from vireo pairs increased because fewer parasitized nests were deserted and parasitized nests on removal plots had a higher probability of success. No cowbird removal study has reported an increase in cowbird productivity in response to trapping. Cowbird nestlings from prairie plots and shrub plots differed in carbon and nitrogen isotope compositions, and 87% of locally produced juvenile cowbirds were classified with nestlings from shrub plots. Thus, the continued expansion of woody plants into tallgrass prairie may result in local increases in cowbird productivity.