Predefinitive Plumage in the Golden Eagle (Aquila chrysaetos): A Signal of Aggression or Submission?

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Explaining variation in brood parasitism rates between potential host species with similar habitat requirements

Host specialization evolved in many parasite-host systems. Evolution and maintenance of host specificity may be influenced by host life-history traits, active host selection by the parasite, and host anti-parasite strategies. The relative importance of these factors is poorly understood in situations that offer parasites a choice between hosts with similar habitat requirements. The common cuckoo Cuculus canorus is a generalist parasite on the species level, but individual females prefer particular host species. In reed beds of the Yellow River Delta, China, two potential hosts with similar nest characteristics, Oriental reed warblers Acrocephalus orientalis and reed parrotbills Paradoxornis heudei, breed in sympatry. We found that warblers were parasitized at much higher rates than parrotbills. Both hosts recognized and rejected non-mimetic model eggs well, indicating that they have been involved in an arms-race with cuckoos. Cuckoo eggs closely resembled warbler eggs, and such eggs were mostly accepted by warblers but rejected by parrotbills. Only warblers recognized adult cuckoos as a specific threat. Both hosts were equally good at raising cuckoo chicks. Low nest density, partial isolation by breeding time, small scale differences in nest and nest site characteristics, and high rejection rates of natural cuckoo eggs are likely cumulatively responsible for the low current parasitism rate in parrotbills. This study emphasizes the importance of integrating the study of general host life-history characteristics and specific anti-parasitism strategies of hosts across all breeding stages to understand the evolution of host specificity.submittedVersionpublishedVersio

CO₂ minimum miscibility pressure determination of pure hydrocarbons in different temperatures using slimtube simulations

As wind energy deployment increases and larger wind-power plants are considered, bird fatalities through collision with moving turbine rotor blades are expected to increase. However, few (cost-) effective deterrent or mitigation measures have so far been developed to reduce the risk of collision. Provision of “passive” visual cues may enhance the visibility of the rotor blades enabling birds to take evasive action in due time. Laboratory experiments have indicated that painting one of three rotor blades black minimizes motion smear (Hodos 2003, Minimization of motion smear: Reducing avian collisions with wind turbines). We tested the hypothesis that painting would increase the visibility of the blades, and that this would reduce fatality rates in situ, at the Smøla wind-power plant in Norway, using a Before–After–Control–Impact approach employing fatality searches. The annual fatality rate was significantly reduced at the turbines with a painted blade by over 70%, relative to the neighboring control (i.e., unpainted) turbines. The treatment had the largest effect on reduction of raptor fatalities; no white-tailed eagle carcasses were recorded after painting. Applying contrast painting to the rotor blades significantly reduced the collision risk for a range of birds. Painting the rotor blades at operational turbines was, however, resource demanding given that they had to be painted while in-place. However, if implemented before construction, this cost will be minimized. It is recommended to repeat this experiment at other sites to ensure that the outcomes are generic at various setting

Methods to quantify avian airspace use in relation to wind energy development

It is likely that there will continue to be a substantial increase in the number of wind turbines as we aim to meet global energy demands through renewable sources. However, these structures can have adverse impacts on airborne wildlife, such as posing a potential collision risk with the turbine structure. A range of methods and technologies have been applied to the collection of bird flight parameters, such as height and speed, to improve the estimation of potential collision compared with traditional visual methods, but these are currently not applied in a consistent and systematic way. To this end, a systematic literature search was conducted to (1) examine the methods and technologies that can be used to provide bird flight data to assess the impact of wind energy developments and (2) provide an updated framework to guide how they might be most usefully applied within the impact assessment process. Four empirical measurement methods were found that improve the estimation of bird flight parameters: radar, telemetry, ornithodolite and LiDAR. These empirical sensor-based tools were typically more often applied in academic peer-reviewed papers than in report-based environmental statements. Where sensor-based tools have been used in the report-based literature, their inconsistent application has resulted in an uncertain regulatory environment for practitioners. Our framework directly incorporates sensor-based methods, together with their limitations and data requirements, from pre-deployment of infrastructure to post-consent monitoring of impacts. This revised approach will help improve the accuracy of estimation of bird flight parameters for ornithological assessment of wind energy. Sensor-based tools may not be the most cost-effective. However, a precedent has been set for wind energy development consent refusal based on ornithological impact assessment, and therefore the cost of collecting accurate and reliable flight data may be balanced favourably against the cost of development consent refusalacceptedVersio

Within-year nest reuse in open-nesting, solitary breeding passerines

During fieldwork at Lake Sic (46°57’N, 23°54’E), Romania, in the summer of 2003, we observed a strange incident of nest reuse in the Great Reed Warbler Acrocephalus arundinaceus. After first laying a complete clutch of five eggs and then ejecting an experimentally added parasite egg together with two of its own eggs, a new clutch was initiated in the same nest. In Tana (70°16’N, 28°19’E), Norway, in June 2003, we observed a similar incident in the Brambling Fringilla montifringilla. In a nest that was completely depredated when 3-4 eggs had been laid, a new clutch was initiated 8-9 days later. This is as far as we know the first time nest reuse has been documented in Great Reed Warblers and Bramblings

Resident bird species track inter-annual variation in spring phenology better than long-distance migrants in a subalpine habitat

The ability to track variation in climate is important for species to persist in a given environment. Lack of responses to both long-term changes and inter-annual variation in climate parameters can result in reduced fitness and population decline. Furthermore, migration strategy can influence the ability to track climatic variation due to the potential to use reliable environmental cues. Here, we studied the temporal relationship between birch leafing and onset of breeding for three bird species with contrasting migration strategies over a 20-year period in a subalpine habitat in Central Norway. We found no temporal change in birch leafing date or breeding onset for the three bird species over the study period. However, we found a statistically significant difference in the ability to track inter-annual variation in birch leafing date between the resident and two long-distance migratory species. The resident great tit Parus major was more capable of initiating egg laying in closer association to variation in birch leafing in early springs, than the long-distance migratory European pied flycatcher Ficedula hypoleuca and common redstart Phoenicurus phoenicurus. Long-distance migrants seem to have been constrained by arrival date or time from arrival to entering the breeding areas, in contrast to resident birds, which might be better able track early initiation of spring in breeding areas by adjusting egg laying date. Our findings highlight the importance of not solely studying directional long-term climatic change, but also pay attention to inter-annual variation

First-Time Migration in Juvenile Common Cuckoos Documented by Satellite Tracking

Being an obligate parasite, juvenile common cuckoos Cuculus canorus are thought to reachtheir African wintering grounds from Palearctic breeding grounds without guidance fromexperienced conspecifics but this has not been documented. We used satellite tracking tostudy naïve migrating common cuckoos. Juvenile cuckoos left breeding sites in Finlandmoving slowly and less consistently directed than adult cuckoos. Migration of the juveniles(N = 5) was initiated later than adults (N = 20), was directed toward the southwest±significantlydifferent from the initial southeast direction of adults±and included strikingly long BalticSea crossings (N = 3). After initial migration of juvenile cuckoos toward Poland, themigration direction changed and proceeded due south, directly toward the winter grounds,as revealed by a single tag transmitting until arrival in Northwest Angola where northernadult cuckoos regularly winter. Compared to adults, the juvenile travelled straighter andfaster, potentially correcting for wind drift along the route. That both migration route and timingdiffered from adults indicates that juvenile cuckoos are able to reach proper winteringgrounds independently, guided only by their innate migration programme.</p

Outcomes of Brood Parasite–Host Interactions Mediated by Egg Matching: Common Cuckoos Cuculus canorus versus Fringilla Finches

Antagonistic species often interact via matching of phenotypes, and interactions between brood parasitic common cuckoos (Cuculus canorus) and their hosts constitute classic examples. The outcome of a parasitic event is often determined by the match between host and cuckoo eggs, giving rise to potentially strong associations between fitness and egg phenotype. Yet, empirical efforts aiming to document and understand the resulting evolutionary outcomes are in short supply.We used avian color space models to analyze patterns of egg color variation within and between the cuckoo and two closely related hosts, the nomadic brambling (Fringilla montifringilla) and the site fidelic chaffinch (F. coelebs). We found that there is pronounced opportunity for disruptive selection on brambling egg coloration. The corresponding cuckoo host race has evolved egg colors that maximize fitness in both sympatric and allopatric brambling populations. By contrast, the chaffinch has a more bimodal egg color distribution consistent with the evolutionary direction predicted for the brambling. Whereas the brambling and its cuckoo host race show little geographical variation in their egg color distributions, the chaffinch's distribution becomes increasingly dissimilar to the brambling's distribution towards the core area of the brambling cuckoo host race.High rates of brambling gene flow is likely to cool down coevolutionary hot spots by cancelling out the selection imposed by a patchily distributed cuckoo host race, thereby promoting a matching equilibrium. By contrast, the site fidelic chaffinch is more likely to respond to selection from adapting cuckoos, resulting in a markedly more bimodal egg color distribution. The geographic variation in the chaffinch's egg color distribution could reflect a historical gradient in parasitism pressure. Finally, marked cuckoo egg polymorphisms are unlikely to evolve in these systems unless the hosts evolve even more exquisite egg recognition capabilities than currently possessed

Coevolutionary adaptations in avian brood parasites and their hosts

Dette prosjektet har satt søkelyset på to problemstillinger knyttet til samevolusjonen mellom parasitt og vert; 1) utvikling av vertstilpasninger som mottrekk mot tilpasninger hos parasitten, med spesiell fokus på eggtilpasninger, og 2) mekanismer som kan forklare den store variasjonen i forsvarsatferd mot kullparasittisme blant ulike verter. 1) Flere gjøk- (Cuculus canorus) stammer eller gentes har utviklet egg som er veldig like vertens egne egg, såkalt eggmimikry, for å vanskeliggjøre vertenes eggavvisning. For å svare på dette har mange verter på sin side gjort det vanskeligere for parasitten ved å utvikle en lavere variasjon i utseende mellom sine egg innen kullet (innenkull-variasjon), og en høyere variasjon mellom egg fra kull til kull (mellomkull-variasjon). Mange nordamerikanske spurvefugler blir benyttet som verter av brunhodetrupialen (Molothrus ater), men denne parasitten har ikke utviklet eggmimikry i forhold til vertseggene. Vi sammenlignet kullvariasjonen mellom spurvefugler i Europa og Nord-Amerika og fant en høyere innenkull-variasjon og en lavere mellomkull-variasjon i eggutseende hos nordamerikanske spurvefugler, selv om forskjellen i innenkull-variasjon mellom kontinentene var mindre enn forventet.Hos europeiske spurvefugler er det i tidligere eksperimenter funnet at det er en sammenheng som forventet mellom avvisningsraten overfor parasittiske ikkemimikry egg og kullvariasjonen i eggutseende. Vi fant at det ikke var noen slik sammenheng hos spurvefugler i Nord-Amerika. Resultatene gir støtte til hypotesen om at parasitter med eggmimikry utøver et betydelig seleksjonstrykk for utvikling av bestemte eggkarakterer hos sine verter. Vi undersøkte om det var noen forskjell i innenkull-variasjon hos avvisere og akseptorer av parasittegg innen bestemte populasjoner av tre europeiske spurvefugler; rørsanger (Acrocephalus scirpaceus), bokfink (Fringilla coelebs) og munk (Sylvia atricapilla). Det ble funnet at det var en signifikant forskjell i innenkull-variasjon i eggutseende mellom avvisere og akseptorer av kunstige ikke-mimikry gjøkegg i en rørsanger-populasjon i Tsjekkia; avviserne hadde en lavere innenkull variasjon enn akseptorer av slike egg. Denne vertspopulasjonen har en intermediær avvisningsrate overfor ikke-mimikry egg. Et tilsvarende forsøk ble utført hos en bokfink-populasjon i Norge og en munk-populasjon i Tsjekkia. Begge artene er meget gode avvisere av ikke-mimikry egg, noe som indikerer at de aller fleste individer er i stand til å avvise slike egg. Vi valgte derfor å benytte egg fra artsfrender i forsøkene med disse artene. I motsetning til hos rørsangeren fant vi at det ikke var noen forskjell i innenkullvariasjon mellom akseptorer og avvisere av fremmede egg hos bokfink og munk. Hos begge artene ble det funnet at avvisningen av fremmede egg i stor grad avhenger av kontrasten (grad av mimikry) mellom egne egg og parasittegget. Dette viser at selv om individene er i stand til å avvise parasittegg, så finnes det kognitive begrensninger som medfører at egg som utseendemessig ligger under en viss terskelverdi med hensyn til likhet med egne egg vil bli akseptert. Det ble ikke funnet noen indikasjoner på at avvisningsatferden var avhengig av vertenes alder eller av kondisjonelle stimuli for noen av de tre artene. Dette kan tyde på at det er en genetisk basert kobling mellom det å kunne gjenkjenne fremmede egg og innenkull-variasjon. 2) Mange vertsarter viser ingen eller kun intermediære avvisningsrater overfor fremmede ikke-mimikry egg. En slik tilsynelatende suboptimal atferd kan skyldes at det er kostnader forbundet med avvisningen som forhindrer evolusjon av perfekt avvisningsatferd. Slike kostnader kan være feilaktig avvisning av egne egg i uparasitterte reir (gjenkjenningsfeil), eller avvisning av egne egg i tillegg til parasittegget i parasitterte reir (avvisningskostnader). Hos gjøkverter, som ved suksessfull gjøkparasittisme har en reproduktiv suksess tilnærmet lik null, vil kun gjenkjenningsfeil være kilde til et potensielt seleksjonstrykk mot utvikling av høy avvisning av fremmede egg. Vi undersøkte om slike kostnader forekommer hos bokfink og munk; to arter som antas å ha blitt benyttet av gjøken tidligere, men som i dag ikke blir regelmessig parasittert. På grunn av at avvisningsatferden opprettholdes i fravær av parasittisme, forventet vi at disse artene begår få gjenkjenningsfeil. Undersøkelsen gav støtte til denne prediksjonen; avvisningskostnader i parasitterte reir var relativt høye, men gjenkjenningsfeil i uparasitterte reir var meget sjeldent forekommende. En hypotese ("spatiell habitat-struktur hypotesen") basert på metapopulasjonsdynamikk og med vekt på karakteristikker vedrørende vertsartenes hekkebiotop ble framsatt for å forklare de store variasjonene i avvisning hos europeiske spurvefugler. Hypotesen bygger på at gjøken benytter de verter som hekker nær utkikkspunkter for parasitten, dvs. nær trær. Arter som hekker både nær og langt fra trær er de beste gjøkvertene, i og med at genflyt fra uparasitterte populasjoner vil forhindre utvikling av perfekt avvisning i parasitterte populasjoner. Arter som alltid hekker nær trær har høye avvisningsrater fordi få eller ingen populasjoner har unnsluppet parasittering, og det har derfor vært sterk seleksjon for utvikling av vertsforsvar. Data for gjøkverter i Europa gav god støtte til hypotesen. Grad av parasitt eggmimikry og parasitteringsrater er høyest hos de vertsarter som kan hekke både langt fra trær og nær trær, noe som tyder på at gjøken har størst suksess hos slike arter.This thesis deals with two topics in the coevolution between brood parasites and their hosts: 1) evolution of host adaptations against parasite egg mimicry, and 2) sources that could explain the considerable variation in rejection behaviour found among various passerines. 1) Several common cuckoo (Cuculus canorus) tribes or gentes in Europe have evolved eggs that are remarkably similar to the host eggs in both size and appearance (i.e. egg mimicry). To counter this adaptation in the parasite, hosts can produce eggs with similar appearance within clutches (low intraclutch variation) as well as eggs with diverging appearance between different clutches (high interclutch variation). Many North American passerines are utilised as hosts by the brown-headed cowbird (Molothrus ater). However, this parasite generally lays non-mimetic eggs. As predicted, we found that European passerines had a lower intraclutch variation and a higher interclutch variation in egg appearance than North American passerines. However, the difference in intraclutch variation between the continents was less than expected. A relationship has previously been found among European passerines between the rejection rate of non-mimetic eggs and clutch variation in egg appearance, and this is thought to reflect the stage in the coevolution between parasite and host. We found no evidence of such patterns among North American species. These results provide support for the hypothesis that specific host clutch variation is a counteradaptation against parasite egg mimicry. We investigated whether there was any difference in clutch variation between acceptors and rejecters of parasitic eggs within populations of three European passerines; reed warblers (Acrocephalus scirpaceus), chaffinches (Fringilla coelebs), and blackcaps (Sylvia atricapilla). In a Czech reed warbler population with an intermediate rejection rate of non-mimetic cuckoo eggs, it was found that rejecters had a statistically significant lower intraclutch variation than acceptors of such eggs. Age or conditional stimuli did not seem to have any influence on the rejectionbehaviour. A similar experiment was carried out in a Norwegian chaffinch population and a Czech blackcap population, which, however, were experimentally parasitised with foreign conspecific eggs because they are both very good rejecters of nonmimetic parasitic eggs. We found no difference in intraclutch variation among acceptors and rejecters of foreign eggs in chaffinches and blackcaps. However, it was found that the rejection of conspecific eggs greatly depends upon the contrast (i.e. mimicry) between the parasitic and their own eggs. It therefore seems that even though individuals have the ability to reject foreign eggs, limitations in their cognitive system entails that parasitic eggs that are too similar to the host eggs will be accepted. We also looked for potential effects of age on rejection behaviour and intraclutch variation, but no relationship between these variables was found. The results indicate that in these three species both rejection behaviour and clutch variation are more or less innate features, and also that there is a genetically based linkage between recognition of odd eggs and intraclutch variation in egg appearance. 2) Many hosts of brood parasites show no or only intermediate rejection rates of foreign non-mimetic eggs. Evolution of proper rejection behaviour could be prevented by costs related to egg rejection. Important in this respect are erroneous rejection of their own eggs in non-parasitised nests (recognition errors) and rejection of their own eggs in addition to the parasitic egg in parasitised clutches (rejection costs). Because successful cuckoo parasitism usually is detrimental to the breeding success of the host, only recognition errors are believed to be important as an opposing selective pressure against proper host defence in cuckoo hosts. We examined whether such costs exist in chaffinches and blackcaps. These species maintain a high rejection rate of foreign eggs, even though they are not currently used as hosts by the cuckoo. We therefore predicted that recognition errors should be absent or at least rare in these species. We found support for this prediction; rejection costs were relatively high but recognition errors were at best rare events. In another investigation, we proposed a hypothesis (the "spatial habitat structure hypothesis") based upon metapopulation dynamics and characteristics concerning host breeding habitats to explain the variation in rejection behaviour found among European passerines. This hypothesis is based upon the fact that the cuckoo, as well as other avian brood parasites, needs access to vantage points in trees to monitor host nests, and thus only species breeding near trees are available as hosts. Our results were very much in accordance with this hypothesis. Species that breed both near and far away from trees are the best cuckoo hosts, because gene flow from non-parasitised populations breeding far from trees will prevent the evolution of proper rejection behaviour in parasitised populations breeding near trees. However, species that always breed near trees have high rejection rates because the majority of the populations have been utilised as hosts, and thus there has been a strong selection for the evolution of host defences. Furthermore, the level of parasite egg mimicry and the level of parasitism was found to be highest among hosts breeding both near and far away from trees, indicating that the cuckoo is most successful when utilising such species as hosts.Paper VI is not included as a paper in this thesis, but is included as the introduction

ERC - Stokke - Understanding the evolution and maintenance of host range in a generalist parasite - the paradox of sympatric host races, 2014

Generalist parasites may expand to new host species, or change the geographic range over which they exploit their hosts, as exemplified by emerging diseases and novel parasites in humans, domesticated plants and animals. Parasite range modifications may bring different host races into contact, facilitating gene flow among them. Sympatric host races of generalist parasites that maintain distinct adaptations to their different hosts represent a great puzzle in evolutionary biology. How can such host races evolve and be maintained in the face of homogenizing gene flow? The aim of the proposed project is to understand the factors that influence evolution and maintenance of host range in a generalist parasite, the common cuckoo Cuculus canorus. In this parasite, individual females specialize on one or a few host species and have evolved specific egg morphs that mimic perfectly those of their hosts. I have compiled a unique database of nearly 40,000 parasitized host clutches found across Europe spanning ca. 200 years, and I will collect phenotypic and genetic data for ca. 2,500 cuckoo eggs stored in European museums. I will analyze spatial and temporal distributions of host races and parasite traits across Europe and determine levels of genetic differentiation and gene flow among host races at both neutral and functional loci. This will allow me to identify factors that determine host range, genetic differentiation and levels of adaptation, and shed light on how host-specific races of parasites evolve and are maintained. Understanding host specificity and host range in a widely distributed generalist parasite will not only have an impact on evolutionary theory, but also with possible ramifications for conservation and veterinary and medical health issues