European whitefish (Coregonus lavaretus) forms reproductively isolated sub-populations with different body sizes in the lakes where northern pike (Esox lucius) exists. It has been hypothesized that the initial reproductive isolation between ecotypes arises through size-assortative mate choice or because small and large individuals arrive to the spawning grounds at different times. To test these hypotheses, I caught and measured both breeding whitefish pairs and singular whitefish on the spawning grounds over the spawning season. There was no correlation between the lengths of males and females in breeding pairs, indicating that mate selection was random with respect to body size. There were no significant differences in body length between sampling dates during the field period in November 2021, suggesting that there is no divergence in spawning time between large and small individuals. Thus, neither of the two hypotheses were supported. Interestingly, males have significantly higher amount of breeding tubercles than females according to my results, which could be a selective factor in mate selection. However, the function of the tubercles is yet poorly understood.

京都大学 / 新制・課程博士 / 博士(理学) / 甲第24457号 / 理博第4956号 / 新制||理||1707(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 中野 隆文, 教授 本川 雅治, 教授 森 哲 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Student Number : 0419128X - MSc Dissertation - School of Animal, Plant and Environmental Sciences - Faculty of Science / Reproductive isolation was investigated in two chromosomally distinct populations of Rhabdomys on the Gauteng highveld. The two populations, Midrand (2n = 48) and Irene (2n = 46), occur 15 km apart, with no known contact or hybrid zone between them. Behavioural experiments, comprising male-female dyadic encounters and female preference tests for same-and different-population male odour, were used to test for premating barriers. Aggression levels were highest in different-population than same-population dyads, and females spent more time with odours of males from their own population than of those of the other population. Breeding and postnatal development studies were conducted to establish postmating barriers. Compared to different-population pairs, reproductive success was markedly reduced in different-population pairings, and the few hybrids that were produced did not breed. My studies indicate that behavioural isolation is well-developed between the Midrand and Irene striped mice, and suggest that the mate recognition system has diverged in allopatry, which would reduce gene flow between the two populations. Such divergence supports the findings of mtDNA studies by other workers who proposed that the two chromosomal forms used in my study represent two subspecies of R. dilectus. Previous studies showed that distant striped mice populations (>900km) displayed behavioural divergence and intermediately located populations (~80km) were behavioural compatible but had hybrid failure; the Irene population was used in both studies. In comparison, the behavioural incompatibility between the closely-located Midrand and Irene populations provides support for the reinforcement of previous postmating isolation seen in the intermediately located populations, particularly since no contact or hybrid zone exists between the two forms. However, I cannot rule out other explanations, such as dissimilar ecological conditions, influencing interfertility.

Três tipos de sons de cortejo foram descritos para machos de tefritídeos: de chamada, na ausência de fêmeas; de aproximação, na proximidade de fêmeas; e de pré-cópula, após a monta das fêmeas pelos machos. Nas espécies do gênero Anastrepha em que foram registrados sons de acasalamento, somente dois tipos estão presentes: de chamada e de précópula. As emissões acústicas de acasalamento de algumas espécies foram caracterizadas quanto a parâmetros como duração e freqüência dos sons, incluindo a espécie nominal Anastrepha fraterculus - importante praga da fruticultura nacional. No entanto, até o presente momento, nenhum estudo teve como foco a diferenciação dentro do complexo A. fraterculus, constituído por diversas espécies crípticas - das quais, três caracterizadas para o Brasil: A. sp. 1 aff. fraterculus, A. sp. 2 aff. fraterculus e A. sp. 3 aff. fraterculus. Este trabalho procurou caracterizar o padrão de emissão acústica de chamada de populações do complexo A. fraterculus (duas de A. sp. 1, uma de A. sp 2 e uma de A. sp. 3) e de A. obliqua, em relação à duração de pulso (PD), intervalo entre pulsos (IPI) e freqüência fundamental (FF), comparando com dados disponíveis na literatura para outras espécies. Os parâmetros acústicos de A. sp. 1 de Serra Negra - SP - não se diferenciaram estatisticamente dos parâmetros de A. sp. 2 de São Sebastião - SP. O padrão de emissão de A. sp. 3 de São Sebastião - SP -, difere do de A. sp. 1 de Serra Negra - SP - e de A. sp. 2 de São Sebastião - SP - no parâmetro PD. Os parâmetros acústicos de A. sp. 1 de São Paulo - SP - foram completamente distintos dos parâmetros das demais populações, inclusive dos de A. sp. 1 de Serra Negra - SP. O padrão de emissão acústica A. obliqua de Águas da Prata - SP - também diferencia-se dos das demais espécies/populações, exceto em relação ao PD de A.. sp. 1 de São Paulo - SP. A relação entre picos de distribuição de FF e de harmônicos, bem como a correlação entre parâmetros acústicos e o comprimento das asas sugerem uma relação entre características físicas das asas e os sons emitidos. Porém, outros fatores também devem estar em atuação, uma vez que a relação entre picos e harmônicos não é observada para todas as populações estudadas e a correlação entre tamanho de asa e valores de parâmetros acústicos é baseada em número limitado de espécies. Não há uma relação entre o valores dos parâmetros acústicos e as relações filogenéticas entre as espécies de moscas-das-frutas. Assim, embora os padrões de emissões acústicas permitam diferenciar as espécies entre si, não é possível utilizá-los para agrupar as espécies quanto a afinidades evolutivas. Dentro dos tefritídeos, a evolução dos tipos de emissão acústica do acasalamento provavelmente envolve uma história bastante complexa com várias homoplasias. / Three courtship songs have been described for tephritid males: calling song, when female flies are absent; approach song, near females; and precopulatory song, after male mounting female. For species of Anastrepha in which mating songs were recorded, only two types are present: calling and precopulatory songs. In these cases mating accoustic emissions were characterized regarding parameters such as duration and sound frequency; including for the nominal species Anastrepha fraterculus - a major fruit crop pest in Brazil. However, no study have focused on differences amid A. fraterculus criptic species complex - including the three species that occur in Brazil: A.. sp. 1 aff. fraterculus, A. sp. 2 aff. fraterculus e A. sp. 3 aff. fraterculus. In this work, the acoustic emission patterns for A. sp. 1, A. sp. 2 A. sp. 3, and for A. obliqua population samples were characterized by comparing pulse duration (PD), interpulse interval (IPI) and fundamental frequency (FF), with data available in literature for other tephritid species. The acoustic parameters of A sp. 1 Serra Negra (SP) do not differ statistically from parameters of A. sp. 2 (São Sebastião, SP). The PD parameter of A. sp. 3 (São Sebastião, SP) differs from A. sp. 1 (Serra Negra, SP) and from A. sp. 2 from São Sebastião (SP). The acoustic parameters of A. sp. 1 from São Paulo (SP) were completely different from parameters of other populations, including A. sp. 1 from Serra Negra (SP). The acoustic emission pattern of A. obliqua from Águas da Prata (SP) is also different from other species/populations, except by the PD of A. sp. 1 from São Paulo (SP). The relationship between FF distribution peaks and sound harmonics as well as the correlation between acoustic parameters and wing length, suggest that the emitted sounds are dependent on the wing physical properties. However, other factors might also influence the sound pattern, because the relation between FF peaks and harmonics is not observed for all population samples, and the correlation between wing length and acoustic parameters were evaluated on few species. There is no relation between acoust parameters values and fruit flies phylogeny. Therefore, it is possible to distinguish between species on the basis of sound parameters, but it is not possible to group them by evolutionary affinities. Among the tephritids, mating acoustic emission patterns probably involves a complex evolutionary history with many homoplasies.

Στόχος της παρούσας εργασίας αποτελεί η διερεύνηση της προσυζευκτικής συμπεριφορικής απομόνωσης στο ερμαφρόδιτο χερσαίο σαλιγκάρι Albinaria caerulea (Pulmonata, Clausiliidae), η πολυπλοκότητα της οποίας, μπορεί να αντανακλά δράση εξελικτικών μηχανισμών επιλογής συντρόφου και κατ' επέκταση ειδογενετικών διεργασιών. Η ποικιλότητα κελυφικής ποίκιλσης είναι έντονη στο είδος αυτό (δύο μορφές, λεία και γραμμωτή, που συνιστούν δυο υποείδη), αλλά δε φαίνεται να συσχετίζεται με αντίστοιχη οικολογική και γενετική διαφοροποίηση. Η συλλογή των ατόμων πραγματοποιήθηκε στη Φολέγανδρο, όπου παρατηρούνται αλλοπάτριοι πληθυσμοί αλλά και ζώνες επαφής μεταξύ των λείων και των γραμμωτών μορφών. Στο εργαστήριο πραγματοποιήθηκαν πειράματα αναπαραγωγικής συμπεριφοράς, όπου καταγράφονταν ο αριθμός κι ο τύπος (ετεροτυπικά-ομοτυπικά) των σχηματιζόμενων ζευγών, καθώς και οι αντίστοιχες ερωτοτροπικές προσπάθειες (φλερτ). Βάσει των συχνοτήτων των σχηματιζόμενων ζευγών υπολογίστηκε ο βαθμός προσυζευκτικής συμπεριφορικής απομόνωσης με τον αμερόληπτο δείκτη ΙPSI. Καταγράφηκαν επίσης και άλλες συμπεριφορικές παράμετροι (αριθμός και τύπος φλερτ, χρόνος ως το πρώτο ενεργητικό φλερτ, χρόνος από την αφύπνιση ως την έναρξη της σύζευξης, διάρκεια σύζευξης), με σκοπό την ανίχνευση λοιπών προσυζευκτικών συμπεριφορικών διαφορών μεταξύ των δυο μορφών. Τέλος, ελέγχθηκε αν τυχόν μορφομετρικές διαφορές (σχήμα/μέγεθος κελύφους) μεταξύ των δύο μορφών, χρησιμοποιούνται στις στρατηγικές επιλογής συντρόφου. Ο βαθμός προσυζευκτικής συμπεριφορικής απομόνωσης βρέθηκε μικρός, καθώς τα άτομα, ως επί το πλείστον, επέλεγαν ετεροτυπικό σύντροφο. Επίσης βρέθηκε μεγαλύτερη προσυζευκτική και συζευκτική δραστηριότητα στις ζώνες επαφής, κάτι που απορρίπτει την υπόθεση περί μηχανισμού ενίσχυσης, .καθώς εάν υπήρχε απομόνωση θα ανιχνεύονταν πρώτα σε αυτές. Όσον αφορά τις μορφομετρικές διαφορές, παρά το ότι τα άτομα διακρίνονται βάσει σχήματος και μεγέθους κελύφους, δεν φαίνεται να χρησιμοποιούν τις διαφορές αυτές στους μηχανισμούς επιλογής συντρόφου. Εντούτοις, οι διαφορές στο μέγεθος κελύφους των δυο μορφών εξομαλύνονται στις ζώνες επαφής, και επομένως δεν υπάρχει απόκλιση χαρακτήρα. Έτσι λοιπόν, ενώ δεν φαίνεται να υπάρχουν ισχυρά προσυζευκτικά φράγματα στο είδος αυτό, περισσότερο φως στις αντίστοιχες εξελικτικές διεργασίες, θα ρίξει η μελέτη μετασυζευκτικών μηχανισμών απομόνωσης, αλλά και παραμέτρων της οικολογίας και της πληθυσμιακής γενετικής τους. / The aim of this study was the investigation of premating behavioural isolation in the hermaphrodite land snail Albinaria caerulea (Pulmonata, Clausiliidae). The snails of this genus exhibit a really complicated premating behaviour, a fact that may reflect evolutionary mechanisms of mate choice, or even speciation forces acting. Shell differentiation is intense, with two patterns (smooth and ribbed) constituting two subspecies, which don’t seem to differ ecologically or genetically. The animals were collected from the island of Folegandros, where they can form contact zones –between animals with smooth and ribbed shell- or the two forms can be isolated. Mate choice experiments were conducted, with variable number of possible mates and the premating behaviour was reported (mating attempts, successful copulations, copulation duration, mating activity), between populations exhibiting the same or different ribbing pattern. To reveal other behavioural differences between the two morphs, we also reported the number and type of courtships, the time from the awakening till the beginning of copulation, the time untill the first active courtship and the duration of copulation. Finally, we examined if possible morphometric differences (shell shape and size) between the two morphs, can be used in the mate choice strategies. The unbiased isolation index IPSI, which based on mate frequencies, suggested that there aren’t strong premating barriers. The intense observed activity (copulations and courtships) in the contact zones agrees with the above results and consequently does not support the initial hypothesis of reinforcement of behavioural barriers in these sites. As far as it concerns the morphometric differences, despite the fact that the morphs are being distinguished, they don’t seem to use them as criterion in the mate choice. Nevertheless, such differences in size, seem to be eliminated in the contact zones, suggesting that there is no character displacement. In conclusion, further investigation of the reproduction of the genus in the basis of postmating isolation mechanisms, the ecology and population genetics of the genus, will reveal the possible action of evolutionary forces.

Les processus d’hybridation et d’introgression occupent une place importante dans l’étude du mécanisme de spéciation, car ils permettent d’analyser les conséquences évolutives des échanges génétiques entre espèces partiellement isolées. Ici, nous nous sommes intéressés à la zone d’hybridation entre les soles S. senegalensis et S. aegyptiaca au niveau des côtes nord-tunisiennes, pour comprendre l’origine de leur diversification. Nous avons premièrement caractérisé les conséquences phénotypiques de l’hybridation sur la variabilité morphologique. Nos résultats montrent que si l’introgression provoque la convergence de certains caractères morphologiques, elle est en revanche à l’origine de transgressions et de distorsions morphologiques sur d’autres traits, pouvant refléter une condition plus faible des génotypes recombinants. Les phénomènes d’incompatibilité génétique associés à une éventuelle contre-sélection des hybrides sont supposés créer une perméabilité différentielle du génome face au flux génique entre espèces. Pour étudier cette semi-perméabilité à l’échelle du génome, nous avons établi un jeu de données de polymorphisme par la méthode RAD-seq. Ceci nous a permis de génotyper 200 individus pour 10 756 marqueurs SNP, qui nous ont permis de caractériser les flux géniques entre ces deux espèces à travers trois approches complémentaires. La première est basée sur une reconstitution démographique de l’histoire des échanges génétiques qui intègre les effets de la semi-perméabilité des génomes. La seconde approche se focalise sur l’évolution spatiale des fréquences alléliques à travers la zone d’hybridation. La dernière méthode, dite des clines génomiques, compare le comportement de chaque locus au patron d’introgression moyen attendu sous l’hypothèse de neutralité. Nos résultats indiquent que S. senegalensis et S. aegyptiaca ont subi une divergence ancienne en allopatrie suivie d’un contact secondaire récent. Seule une faible proportion du génome parvient à introgresser de manière asymétrique dans la zone hybride qui en résulte, selon une grande diversité de patrons d’introgression dont nous discutons les origines possibles. / Hybridisation and introgression processes have an important place in the study of speciation as they allow to analyse the evolutionary consequences of genetic exchanges between partially isolated species. Here we are interested in the hybrid zone resulting from the contact between the soles S. aegyptiaca and S. senegalensis along the North Tunisian coast. First, we studied the consequences of hybridisation on phenotypic variation. This allowed us to evidence that phenotypic convergence in hybrids was accompanied by phenotypic transgression and morphological distortions for certain traits that seem to reflect a reduced condition of hybrids. Possible genetic incompatibilities between species should be responsible for the differential permeability of the genome to gene flow, thereby creating a semi-permeable barrier. To study this barrier at the genome scale, we have produced a polymorphism dataset using the RAD-seq method. This allowed us to genotype 200 individuals at 10,756 SNP markers and to characterise genomic patterns of gene flow between the two species through three complementary approaches. The first is based on a reconstruction of the demographic history of the genetic and exchanges that incorporates the effects of the semi-permeability to gene flow. The second approach focuses on the spatial evolution of allele frequencies across the hybrid zone. The last method, called genomic clines, compares the behaviour of each locus to the average introgression pattern expected under the hypothesis of neutrality. Our results indicate that S. aegyptiaca and S. senegalensis underwent ancient divergence in allopatry followed by a recent secondary contact. Only a small proportion of the genome can asymmetrically introgress across the hybrid zone, resulting in a variety of introgression patterns of which we discuss the possible origins.

Made available in DSpace on 2014-06-11T19:29:15Z (GMT). No. of bitstreams: 0 Previous issue date: 2011-02-21Bitstream added on 2014-06-13T20:19:16Z : No. of bitstreams: 1 salviano_mb_me_jabo.pdf: 2171806 bytes, checksum: 708a5f60d73e3222397411ea4a47e2c5 (MD5) / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) / A espécie Mazama americana tem sido bastante estudada nas áreas de ecologia, genética e reprodução devido às recentes dúvidas quanto à validade de seu táxon. Estudos citogenéticos e moleculares demonstram que atualmente este táxon é composto de uma espécie que apresenta até seis citótipos diferentes. Instituições voltadas à conservação classificam este táxon como “Dados insuficientes”, devido às incertezas taxonômicas. Este trabalho teve como objetivo avaliar a fertilidade dos animais resultantes de cruzamentos intra e intercitótipos para inferir sobre a existência de isolamento reprodutivo entre os citótipos e conclusões acerca da existência de espécies novas. Foram avaliados o sêmen, histologia testicular, perfil de metabólitos fecais de testosterona e a meiose de sete animais, sendo dois puros (cruzamento intracitótipo), três híbridos de mesma linhagem evolutiva e dois híbridos de linhagens evolutivas diferentes. Os resultados mostram que os animais puros tiveram melhores desempenhos reprodutivos seguidos pelos animais híbridos entre citótipos de mesmas linhagens evolutivas, que foram sugestivos de subfertilidade. Os animais de linhagens evolutivas diferentes apesar de apresentarem atividade estereidogênica foram azoospérmicos o que nos permite concluir que dentro do táxon existem pelo menos duas espécies diferentes com possibilidade de duas subespécies dentro de cada ramo filogenético. Estes resultados implicam na necessidade de novas caracterizações taxonômicas, inclusive para que sejam estruturados planos de ação para conservação de espécies que pudessem estar correndo risco de extinção / Mazama americana (Mammalia, Cervidae), has been widely studied in ecologic, genetic and reproductive areas given recent doubts about its taxonomic validity. Cytogenetic and molecular studies showed that currently this taxon is composed by a species with up to six different cytotypes. The World Conservation Union IUCN classified this taxon as “data deficient” given the taxonomic uncertain of this species. This research aimed to evaluate the fertility of of intra and intercytotypic breeding products in order to conclude about the occurrence of novel species. Semen analysis, testicular histology, fecal testosterone metabolites and meiosis of seven breeding products, being two of them pure (intracytotipic breeding), three hybrids from the same evolutive lineage and two hybrids from different evolutive lineages. Results show that pure products have better reproductive performances, being the hybrids from the same evolutive lineage suggestive of subfertility. The hybrid products from different evolutive lineages were azoospermics, despite their steroid activity. These results show the occurrence of almost two different species and the possibility of almost four subspecies. It implies the necessity of new taxonomic review, which will aid in the design of action plans for the conservation of endangered species

Orientador: José Maurício Barbanti Duarte / Banca: Luiz Renato de França / Banca: Vera Fernanda Martins Hossepian de Lima / Resumo: A espécie Mazama americana tem sido bastante estudada nas áreas de ecologia, genética e reprodução devido às recentes dúvidas quanto à validade de seu táxon. Estudos citogenéticos e moleculares demonstram que atualmente este táxon é composto de uma espécie que apresenta até seis citótipos diferentes. Instituições voltadas à conservação classificam este táxon como "Dados insuficientes", devido às incertezas taxonômicas. Este trabalho teve como objetivo avaliar a fertilidade dos animais resultantes de cruzamentos intra e intercitótipos para inferir sobre a existência de isolamento reprodutivo entre os citótipos e conclusões acerca da existência de espécies novas. Foram avaliados o sêmen, histologia testicular, perfil de metabólitos fecais de testosterona e a meiose de sete animais, sendo dois puros (cruzamento intracitótipo), três híbridos de mesma linhagem evolutiva e dois híbridos de linhagens evolutivas diferentes. Os resultados mostram que os animais puros tiveram melhores desempenhos reprodutivos seguidos pelos animais híbridos entre citótipos de mesmas linhagens evolutivas, que foram sugestivos de subfertilidade. Os animais de linhagens evolutivas diferentes apesar de apresentarem atividade estereidogênica foram azoospérmicos o que nos permite concluir que dentro do táxon existem pelo menos duas espécies diferentes com possibilidade de duas subespécies dentro de cada ramo filogenético. Estes resultados implicam na necessidade de novas caracterizações taxonômicas, inclusive para que sejam estruturados planos de ação para conservação de espécies que pudessem estar correndo risco de extinção / Abstract: Mazama americana (Mammalia, Cervidae), has been widely studied in ecologic, genetic and reproductive areas given recent doubts about its taxonomic validity. Cytogenetic and molecular studies showed that currently this taxon is composed by a species with up to six different cytotypes. The World Conservation Union IUCN classified this taxon as "data deficient" given the taxonomic uncertain of this species. This research aimed to evaluate the fertility of of intra and intercytotypic breeding products in order to conclude about the occurrence of novel species. Semen analysis, testicular histology, fecal testosterone metabolites and meiosis of seven breeding products, being two of them pure (intracytotipic breeding), three hybrids from the same evolutive lineage and two hybrids from different evolutive lineages. Results show that pure products have better reproductive performances, being the hybrids from the same evolutive lineage suggestive of subfertility. The hybrid products from different evolutive lineages were azoospermics, despite their steroid activity. These results show the occurrence of almost two different species and the possibility of almost four subspecies. It implies the necessity of new taxonomic review, which will aid in the design of action plans for the conservation of endangered species / Mestre

Un objectif central en biologie est de comprendre la relation entre le génotype et le phénotype. Mes travaux de thèse s’inscrivent dans cette thématique et focalisent sur les populations naturelles d’une même espèce, en utilisant la levure Saccharomyces cerevisiae comme système d’étude. Dans un premier temps, je me suis focalisée sur l’effet des mutations présentes dans différents isolats naturels sur l’apparition de l’isolement reproductif, un processus engendrant la perte de la viabilité de la descendance lors du croisement. Dans un second temps, en plus des phénotypes sévères tels que la létalité de la descendance, je me suis aussi intéressée à la caractérisation de la complexité génétique d’un ensemble de traits quantitatifs tels que la croissance sur différents conditions de culture, afin d’avoir une vision globale du nombre de gènes impliqués et de leur mode d'interaction qui sous-tend la variation phénotypique au sein d’une même espèce. / Elucidating the genetic origin of phenotypic diversity among individuals within the same species is essential to understand evolution. Using the yeast Saccharomyces cerevisiae, we showed that reproductive isolation could readily segregate at the intraspecific level, which is governed by various molecular mechanisms ranging from large-scale chromosomal changes to incompatible epistatic genetic interactions. Compared to reproductive isolation, other phenotypes such as monogenic Mendelian traits are thought to be simple in terms of their phenotypic penetrance and genetic constitution. However, our survey showed that the expressivity of monogenic mutations and hence the inheritance pattern of a Mendelian trait could also depend on parental combinations, transitioning from simple to complex. Our studies unveiled the multiplicity and complexity of the genetic origin of phenotypes within a population, from the origin of reproductive isolation to the hidden complexity of Mendelian traits.

A critical step in ecological speciation is the development of reproductive isolation. The processes leading up to reproductive isolation are difficult to study since they often occur over very long time periods. Populations of European whitefish (Coregonus lavaretus) has gone through reticulate divergent evolution in many Scandinavian lakes during the last 10 000 years. Some populations even exhibit morphological and genetic divergence that involves adaptations to different niches during the first 100-200 years after introduction. This observed rapid diversification into different ecological niches makes whitefish a useful model species for studying ecological speciation and early population divergence. By assessing divergence for three traits in recently introduced whitefish populations, this study aims to elucidate the processes that lead to adaptive phenotypic divergence and reproductive isolation. Whitefish populations of known age (71-170 years) were sampled on their spawning grounds and characterized with respect to, 1) length of spawning season, 2) temporal segregation based on body size, and 3) temporal segregation based on the number of gill rakers. I found that the length of the spawning season and body size differences between early and late spawners increased with population age. No such trend was observed for differences in gill rakers between early and late spawners, but significant divergence within some of the older populations was detected. I conclude that these young whitefish populations have taken the first steps toward reproductive isolation between ecotypes that differs in body size (a highly plastic trait) and gill raker numbers (a trait under strong genetic control).