Genomic organization of the major histocompatibility complex (MHC) can profoundly influence gene function and multigene family evolution. Situated at the interface of individual genetic variation and the adaptive immune response, MHC class I and II loci are intensively studied for disease associations and used as markers of adaptive genetic variation in evolutionary ecology research. Genomic sequence of MHC-containing cosmid clones from the red-billed gull (Larus scopulinus, Charadriiformes: shorebirds, gulls, and allies) was obtained for comparative analysis of avian MHC evolution. MHCI polymorphism was further investigated using cDNA library screening and locus-specific genotyping protocols. This first information regarding MHC organization and MHCI variation in charadriiforms suggests a complex evolutionary history to MHC architecture in birds. Duplication of MHCIIα loci in tandem MHCIIα/β pairs and their proximity to MHC-region gene COL11A2 are similar to arrangements in nonavian vertebrates, and contrast with the “minimal essential” MHC of the chicken (Gallus gallus, Galliformes: gamebirds). MHCI–TAP2 organization is shared with Galloanserae (gamebirds + waterfowl), as is a proposed major classical function for this MHCI gene. In contrast, the placement of MHCI genes adjacent to sequence from chromosomes 3, 5, and 22 of the chicken and zebra finch (Taeniopygia guttata, Passeriformes: perching birds) indicates interchromosomal rearrangements in birds and the possible genomic dispersal of nonclassical MHCI genes in the red-billed gull. Screening for avian malaria, genetic parentage tests, and field data from red-billed gulls at Kaikoura Peninsula, New Zealand were combined with MHCI genotypes to investigate relationships with disease and reproduction. Plasmodium infection was confirmed in red-billed gulls, and breeding condition was negatively associated with malarial infection and positively related to variation at the putative major MHCI locus. A low rate of extrapair paternity was identified across thirteen breeding seasons. Partners without extrapair young (EPY) had greater MHCI dissimilarity than was expected by chance, whereas lower individual MHCI variation and elevated hatching failure existed for pairs with EPY. In addition to contributing to studies of MHC evolution, sexual selection, and disease dynamics in the New Zealand avifauna, this research will facilitate studies of MHC genes in related charadriiforms, many of which are of conservation concern.

Genomic organization of the major histocompatibility complex (MHC) can profoundly influence gene function and multigene family evolution. Situated at the interface of individual genetic variation and the adaptive immune response, MHC class I and II loci are intensively studied for disease associations and used as markers of adaptive genetic variation in evolutionary ecology research. Genomic sequence of MHC-containing cosmid clones from the red-billed gull (Larus scopulinus, Charadriiformes: shorebirds, gulls, and allies) was obtained for comparative analysis of avian MHC evolution. MHCI polymorphism was further investigated using cDNA library screening and locus-specific genotyping protocols. This first information regarding MHC organization and MHCI variation in charadriiforms suggests a complex evolutionary history to MHC architecture in birds. Duplication of MHCIIα loci in tandem MHCIIα/β pairs and their proximity to MHC-region gene COL11A2 are similar to arrangements in nonavian vertebrates, and contrast with the “minimal essential” MHC of the chicken (Gallus gallus, Galliformes: gamebirds). MHCI–TAP2 organization is shared with Galloanserae (gamebirds + waterfowl), as is a proposed major classical function for this MHCI gene. In contrast, the placement of MHCI genes adjacent to sequence from chromosomes 3, 5, and 22 of the chicken and zebra finch (Taeniopygia guttata, Passeriformes: perching birds) indicates interchromosomal rearrangements in birds and the possible genomic dispersal of nonclassical MHCI genes in the red-billed gull. Screening for avian malaria, genetic parentage tests, and field data from red-billed gulls at Kaikoura Peninsula, New Zealand were combined with MHCI genotypes to investigate relationships with disease and reproduction. Plasmodium infection was confirmed in red-billed gulls, and breeding condition was negatively associated with malarial infection and positively related to variation at the putative major MHCI locus. A low rate of extrapair paternity was identified across thirteen breeding seasons. Partners without extrapair young (EPY) had greater MHCI dissimilarity than was expected by chance, whereas lower individual MHCI variation and elevated hatching failure existed for pairs with EPY. In addition to contributing to studies of MHC evolution, sexual selection, and disease dynamics in the New Zealand avifauna, this research will facilitate studies of MHC genes in related charadriiforms, many of which are of conservation concern.

As a contribution to the existing knowledge of grooming in primates five and a half years of grooming data were examined from a group of free-living chimpanzees (Pan troglodytes) in the Budongo Forest, Uganda, to investigate various functional significances of grooming behaviour within the context of social reinforcement. The fission–fusion social structure of chimpanzees results in group members not moving around as a single unit, but forming temporary units as the need arises. This reduces opportunities for individuals to groom others and therefore, based on time and association constraints alone, grooming was as expected found to be unevenly distributed among group members. Grooming patterns found among this group of chimpanzees were comparable to those observed in other free-living populations with variations possibly being attributed to resource base, population numbers and differences in age-sex class composition. One of the suggested social benefits of grooming is that it is used to enhance reproductive success, either by allowing males to enhance their proximity to oestrous females, or by influencing female choice through the development of affiliative relationships with males. Grooming was found to increase between males and females, whilst females displayed sexual receptivity through the presence of anogenital swellings and grooming may be a strategy used by males to increase their access to copulation opportunities, whereas females may use grooming to increase protection from harassment by less preferred males during swollen periods and also increase the likelihood of copulation with preferred partners. Based on the availability of oestrous females, copulations between males and adult females occurred significantly less frequently than expected, whereas copulations between males and subadult females occurred significantly more frequently than expected. Overall a positive correlation was found between grooming of females by males and frequency of copulations. Due to concerns regarding the validity of different sampling methods, scan-focal and ad libitum sampling methods were compared to establish if results from different sampling methods were similar. Results from the scan-focal and ad libitum sampling methods had very few discrepancies, and it is suggested that ad libitum sampling methods which record behaviour types whenever they occur, may be more beneficial for species which don’t move around as a single unit and live in environments where visibility is reduced, therefore increasing the possibility of recording individuals or behaviours that are observed infrequently. Scan-focal sampling may be more beneficial in studying species which move around together in habitats which are conducive to greater visibility, therefore allowing all or most group members to be observed simultaneously. / Thesis (DPhil)--University of Pretoria, 2009. / Zoology and Entomology / unrestricted

Cardiocondyla elegans est une fourmi méditerranéenne que l'on retrouve sur les bords de Loire. Cette espèce possède quelques particularités, comme par exemple avoir des mâles aptères. L'objectif de cette thèse était de comprendre sa biologie, ses stratégies de reproduction ainsi que les caractéristiques de son environnement. Pour cela, nous avons utilisé une approche multidisciplinaire, menant des études de pédologie, d'écologie et de botanique (caractérisation de l'habitat et de l'environnement), ainsi que de génétique et d'éthologie (structure génétique des nids et des populations, détermination du génotype et comportement des sexués).<br />Après avoir confirmé génétiquement la monogynie de cette espèce (une seule reine entourée d'environ 200 ouvrières), nous nous sommes attachés à analyser l'habitat de C. elegans. Le nid est creusé dans le sol jusqu'à 40 cm de profondeur. Il est composé, comme celui de nombreuses autres espèces, d'un conduit vertical reliant une dizaine de chambres superposées. En milieu ligérien, C. elegans est retrouvée uniquement sur les grèves en fixation. Nous avons montré que la composition des sédiments de ces grèves était importante pour la survie de cette fourmi. Durant l'été, elle permet dans les chambres les plus profondes, de conserver une température constante inférieure à 30°C avec de très faibles variations quotidiennes. Durant les crues, les sédiments composés de moins de 60% de sable, ne sont que très peu entraînés par le courant (grèves « fixées »). De plus ils permettent, lors de la remontée de la nappe alluviale, la création de poches d'air nécessaires à la survie des fourmis.<br />L'étude de la répartition spatiale des nids au cours des années nous a révélé que 40% des nids disparaissent d'une année sur l'autre suite aux conditions hivernales. Cependant, suffisamment d'individus sexués, en l'occurrence des femelles fécondées, survivent pour fonder de nouveaux nids. La population étudiée de C. elegans sur les bords de Loire montre une forte densité avoisinant 1 nid/m2. Les grèves fixées étant des environnements morcelés, les nids entrent en compétition pour l'espace et se distribuent de façon régulière. Ainsi, moins de 1% des femelles réussissent à fonder de nouveaux nids.<br />De Juillet à Septembre, les nids matures produisent des individus sexués. Alors que les femelles sont ailées, C. elegans possède une particularité par rapport aux autres fourmis : elle ne produit que des mâles ergatoïdes (sans ailes) tolérants entre eux. Nous avons déterminé que lors de la période de reproduction, les nids contenaient en moyenne 5,3 mâles ergatoïdes et 76,6 femelles ailées. La reine est généralement fécondée par plusieurs mâles. Ainsi les individus du nid sont issus de la même mère mais peuvent avoir des pères différents (en moyenne 4,5 fratries par nid).<br />En analysant la structure génétique des populations nous pouvons dire que 30% des accouplements impliquent des individus non apparentés. Ce fait est du à la présence de nombreux sexués étrangers à l'intérieur des nids. La présence d'individus étrangers dans des colonies monogynes est singulière et nous a amené à nous interroger sur les stratégies de reproduction de C. elegans. Des tests comportementaux nous ont permis de mettre en évidence que les mâles étaient toujours acceptés lorsqu'ils tentaient de pénétrer dans une nouvelle colonie contrairement aux femelles ailées qui sont systématiquement attaquées. La présence de femelles ailées dans des nids étrangers s'explique par leur transport par des ouvrières, un comportement que nous avons observé sur le terrain en période de reproduction. Ces échanges de sexués pourraient permettre de diminuer la consanguinité à l'intérieur de la population, évitant ainsi la production de mâles diploïdes (stériles) et procurer une meilleure résistance de la population en cas de changements environnementaux.<br />L'étude de cette fourmi nous a permis de mettre en évidence une stratégie de reproduction originale, décrite ici pour la première fois. Notre étude révèle également que C. elegans est la seule espèce de fourmis à être adaptée aux conditions environnementales des grèves en fixation. La seule présence de cet organisme, au même titre que certaines espèces végétales, permet de caractériser une étape de l'évolution morphologique des chenaux secondaires de la Loire.