<p>Speciation is the process in which reproductive barriers evolve between populations. In this thesis I examine how sexual signals contribute to the maintenance, reinforcement or breakdown of reproductive barriers.</p><p>Male pied flycatchers (<i>Ficedula hypoleuca</i>) and collared flycatchers (<i>F. albicollis</i>) differ in song and plumage traits. However, where the two species coexist, several pied flycatchers sing a song resembling the collared flycatcher (mixed song). Mixed song is not caused by introgression from the collared flycatcher but is due to heterospecific copying. Mixed song provokes aggressive behaviour in collared flycatcher males and leads to heterospecific pairing and maladaptive hybridization. </p><p>The species differences in song were found to be larger in an old than a young hybrid zone. This was due to a reduction in the frequency of mixed song in the pied flycatcher and a divergence in the song of the collared flycatcher. Apparently, mixed song causes maladaptive hybridization, which over time leads to reinforcement of reproductive barriers by a song divergence.</p><p>Previous studies have shown that a character displacement in male plumage traits reinforces species barriers. Hence both plumage and song divergence reduce the incidence of hybridization. The evolution of male plumage traits has been so rapid, or selection has been so strong that rapidly evolving molecular markers are unable to trace it.</p><p>Hybrid females mate with a male of the same species as their father. Previous studies have shown that females use male plumage traits controlled by genes linked to the sex chromosomes (the Z) in species recognition. An association between preference and a sex-linked trait through the paternal line may render reinforcement of reproductive barriers more likely.</p><p>In conclusion, sexual signals are affected by species interactions that cause breakdown or reinforcement of reproductive barriers.</p>

Speciation is the process in which reproductive barriers evolve between populations. In this thesis I examine how sexual signals contribute to the maintenance, reinforcement or breakdown of reproductive barriers. Male pied flycatchers (Ficedula hypoleuca) and collared flycatchers (F. albicollis) differ in song and plumage traits. However, where the two species coexist, several pied flycatchers sing a song resembling the collared flycatcher (mixed song). Mixed song is not caused by introgression from the collared flycatcher but is due to heterospecific copying. Mixed song provokes aggressive behaviour in collared flycatcher males and leads to heterospecific pairing and maladaptive hybridization. The species differences in song were found to be larger in an old than a young hybrid zone. This was due to a reduction in the frequency of mixed song in the pied flycatcher and a divergence in the song of the collared flycatcher. Apparently, mixed song causes maladaptive hybridization, which over time leads to reinforcement of reproductive barriers by a song divergence. Previous studies have shown that a character displacement in male plumage traits reinforces species barriers. Hence both plumage and song divergence reduce the incidence of hybridization. The evolution of male plumage traits has been so rapid, or selection has been so strong that rapidly evolving molecular markers are unable to trace it. Hybrid females mate with a male of the same species as their father. Previous studies have shown that females use male plumage traits controlled by genes linked to the sex chromosomes (the Z) in species recognition. An association between preference and a sex-linked trait through the paternal line may render reinforcement of reproductive barriers more likely. In conclusion, sexual signals are affected by species interactions that cause breakdown or reinforcement of reproductive barriers.

Speciation can be viewed as the formation of reproductive barriers between different populations. This thesis investigates patterns of natural and sexual selection shaping reproductive barriers between two hybridizing flycatchers (i.e. collared – and pied flycatchers). Behaviorally driven sexual isolation depends on both the availability of conspecific mates and on discrimination ability of individuals. My results demonstrate that these two factors may also interact. Discrimination abilities may change in response to the relative frequency of two interbreeding species. The underlying reason appears to be that male pied flycatchers have a song that incorporates more elements of the song characteristics of male collared flycatchers into their own song repertoires when occurring in areas inhabited predominantly by collared flycatchers. I investigated selection pressures acting on hybrids. In migratory species, hybrid fitness might be reduced as a consequence of intermediate suboptimal migration routes (extrinsic post zygotic isolation). Comparison of stable isotope signatures of revealed that parental species have separate wintering grounds, but hybrids appear to winter at the same location as pied flycatchers. A possible dominance effect in the inheritance of migration direction may hence reduce this potential cost. This interpretation is supported by the absence of a reduction in juvenile to adult survival of hybrids. By further comparing male hybrid fitness to that of the parental species, using lifehistory data, I demonstrate that hybrid males experience a moderate reduction in fitness (mainly through a sexually selected disadvantage). Sexual selection acting on male hybrids can play a major role in the speciation process because when the same characters affect assortative mating as well as hybrid fitness, reinforcement of reproductive barriers becomes more likely. Even when reproductive isolation is completed- the fate of newly formed species may be uncertain since they may strongly compete for ecological space. Long-term persistence of ecologically similar, species requires that there are spatial or temporal variation in their relative fitness. The growth of nestling pied flycatchers is less affected by harsh environmental conditions. We suggest that a regional co-existence of the two flycatcher species is due to a lifehistory trade-off between interference competitive ability and robustness to a harsh conditions. Overall, the studies in this thesis reveal the complexity of the interactions between mate choice and competition in shaping sexual signals. Furthermore, it suggests that natural selection is moderate on hybrid males and that sexual selection may have strong implications for the maintenance of species integrity.

Abstract Sexual ornaments have developed in a very wide variety of animal taxa to increase fitness by improving mating success. How increased mating rate improves fitness is obvious in the case of most males: each mating provides more offspring. Whether more matings benefit females, whose fecundity is limited by resources and not mates, is unclear. Mate choice is linked to ornamentation, when individuals of one sex choose who to mate with based on the ornamentation in members of the other sex. Ornaments may work as a basis for mate choice due to conveying information about the quality of the mate. Mate choice may then lead to intrasexual competition for mates in the chosen sex as animals try to outcompete their neighbours and attract more mates. My aim in this thesis is to study the purpose and costs of female ornamentation as well as female competition for males. The main questions revolve around understanding what information female ornaments provide about the bearer and how males choose between females. I also examine how females compete against each other and what the costs of being ornamented are for a female. To study these questions I use the common glow-worm (Lampyris noctiluca Linnaeus, Lampyridae), whose females glow at night to attract flying males. According to my results, a female’s glowing provides accurate information about fecundity and males base their choice of mate on the intensity of female glowing. I showed that the perceived strength of an ornament in comparison to others close by is more important than the actual strength of an ornament. In the glow-worm, the perceived strength of an ornament depends on distance to the observer, making the system open to exploitation. A mathematical model I developed suggested that dull females could outcompete brighter ones by choosing optimal locations, and experimentation showed this to be the case. Finally, my results show that glow-worm females lose eggs each day they remain unmated. This suggests that the evolution of female ornamentation may have been driven in part by the need to find a mate as soon as possible after eclosion. Which in turn means that female and male ornamentation may have fundamentally different purposes: for females mating soon may be important, where for males the total number of offspring fertilized is critical. / Tiivistelmä Seksuaalisia ornamentteja on kehittynyt hyvin laajaan kirjoon eläinlajeja parantamaan yksilöiden kelpoisuutta nostamalla niiden parittelutodennäköisyyttä. Koirailla parittelutodennäköisyyden ja kelpoisuuden välinen yhteys on yleensä selvä: jokainen parittelu nostaa jälkeläismäärää. Naarailla yhteys ei ole yhtä selvä, sillä niillä jälkeläismäärä yleensä riippuu resurssien saatavuudesta eikä kumppanien määrästä. Parinvalinta on kiinteästi yhteydessä ornamentaatioon: Yhden sukupuolen yksilöt päättävät kenen kanssa paritella toisen sukupuolen yksilöiden ornamenttien laadun perusteella. Ornamentit toimivat valinnan perusteena välittämällä tietoa parin laadusta. Parinvalinta voi johtaa yksilöiden väliseen kilpailuun kumppaneista valitun sukupuolen sisällä. Tässä väitöskirjassa tarkoituksenani on tutkia naaraiden ornamenttien tarkoitusta ja kustannuksia sekä tutkia miten naaraat voivat kilpailla toisiaan vastaan koiraista. Pääkysymykset liittyvät ornamenttien välittämään informaatioon ja siihen miten koiraat valitsevat kumppaninsa. Lisäksi tarkastelen miten naaraat voivat kilpailla keskenään ja mitä kustannuksia ornamenteista on naaraille. Kysymysten selvittämiseksi käytän tutkimuslajina kiiltomatoa (Lampyris noctiluca Linnaeus, Lampyridae), lajia jonka naaraat loistavat öisin houkutellakseen lentäviä koiraita. Tulosteni mukaan kiiltomatonaaraan loiste välittää tarkkaa informaatiota niiden munamäärästä ja koiraat osaavat valita parinsa tämän perusteella. Osoitin, että ornamentin havaittu vahvuus verrattuna ympärillä oleviin on tärkeämpää kuin ornamentin todellinen vahvuus. Kiiltomadolla havaittu kirkkaus riippuu etäisyydestä havaitsijaan, mikä tekee signallointijärjestelmästä avoimen väärinkäytölle. Laatimani matemaattisen mallin mukaan himmeät naaraat voisivat päihittää kilpailussa kirkkaampia valitsemalla parempia loistamispaikkoja. Käytännön kokeet osoittivat tämän pitävän paikkansa. Viimeinen tulokseni oli, että kiiltomadolla naaras menettää osan munistaan joka päivä jonka se joutuu odottamaan parittelua. Tästä voi päätellä, että naaraan ornamentin evoluutiota on voinut osaltaan ajaa tarve paritella mahdollisimman pian aikuistumisen jälkeen. Naaraiden ja koiraiden ornamenttien syyt voivat olla hyvin erilaiset: naarailla parittelu mahdollisimman pian voi olla tärkeintä, kun koiraalla parittelukumppanien määrä on tärkein.