Hygienic behaviour in the honeybee (Apis mellifera) has been shown to be an effective control mechanism against brood diseases such as chalkbrood and AFB. Chalkbrood has proven to be problematic for the Australian honey industry since it was identified here in 1993. Hygienic behaviour is a much studied trait. Rothenbuhler investigated the genetic basis of hygienic behaviour, proposing a two-gene model to explain the uncapping and removal of dead brood. His elegant experiment remains the textbook example of a behavioural genetic study. Although this model has been challenged, it is still generally agreed that a small number of unlinked genes produce a large effect on hygienic behaviour, that hygienic alleles are recessive and are inherited in a Mendelian manner. Experimental backcross colonies were produced from an inbred hygienic line and an inbred non-hygienic line, both provided by Dr. Marla Spivak, University of Minnesota. These backcross colonies were assessed for hygienic behaviour using a standard assay. Statistical analyses of the field data indicated that the genetic basis of the trait was more complex than either the simple Mendelian and widely accepted two-gene or three-gene models that have been proposed previously. Molecular techniques, linkage mapping and QTL analysis then were employed to determine how many loci directly influence hygienic behaviour and the relative level of influence and location of each locus within the genome of A. mellifera. Full multipoint linkage analysis by Mapmaker v3.0 software produced a new genetic map of the honeybee comprised of 358 marker loci ordered over 25 linkage groups spanning a total distance of 3406.2 cM. The average distance between each marker was 9.5 cM. QTL analysis of the experimental data identified seven putative genetic markers associated with hygienic behaviour. QTLs located on linkage groups 2, 4, 6 and 22 were detected for both overall hygienic behaviour and uncapping behaviour only. Individually, each QTL is of relatively small effect with each explaining only 9% � 15% of the variance in hygienic levels observed. Collectively, the putative QTLs identified here explain 79.4% of the observed variance in the expression of hygienic behaviour. These results indicate that there are many genes of low to moderate effect rather than few genes of large effect involved in this complex behavioural trait. This is typical of inherited quantitative traits which do not exhibit Mendelian phenotypic ratios. DNA extracted from the brood samples taken during testing of commercial stock, and from individual bees identified as either highly hygienic or non-hygienic in a reciprocal backcross experiment, were screened with the candidate markers associated with putative QTLs to test their diagnostic power. Unfortunately, none have produced reliably diagnostic DNA profiles. As we have now shown that hygienic behaviour is a polygenic, quantitative trait, simple diagnostic markers for Rothenbuhler's 'uncapping' and 'removal' genes are unlikely to be achieved. Our results show that the most likely way to improve disease resistance in Australian stock is via traditional methods of recurrent selection. The project was responsible for the importation of new genetic material into Australia from the United States. This hygienic stock has been well received by industry, has been widely disseminated, and incorporated into local breeding programs. We hope that it has lead to a general improvement in the level of disease resistance in Australian commercial bees.

Hygienic behaviour in the honeybee (Apis mellifera) has been shown to be an effective control mechanism against brood diseases such as chalkbrood and AFB. Chalkbrood has proven to be problematic for the Australian honey industry since it was identified here in 1993. Hygienic behaviour is a much studied trait. Rothenbuhler investigated the genetic basis of hygienic behaviour, proposing a two-gene model to explain the uncapping and removal of dead brood. His elegant experiment remains the textbook example of a behavioural genetic study. Although this model has been challenged, it is still generally agreed that a small number of unlinked genes produce a large effect on hygienic behaviour, that hygienic alleles are recessive and are inherited in a Mendelian manner. Experimental backcross colonies were produced from an inbred hygienic line and an inbred non-hygienic line, both provided by Dr. Marla Spivak, University of Minnesota. These backcross colonies were assessed for hygienic behaviour using a standard assay. Statistical analyses of the field data indicated that the genetic basis of the trait was more complex than either the simple Mendelian and widely accepted two-gene or three-gene models that have been proposed previously. Molecular techniques, linkage mapping and QTL analysis then were employed to determine how many loci directly influence hygienic behaviour and the relative level of influence and location of each locus within the genome of A. mellifera. Full multipoint linkage analysis by Mapmaker v3.0 software produced a new genetic map of the honeybee comprised of 358 marker loci ordered over 25 linkage groups spanning a total distance of 3406.2 cM. The average distance between each marker was 9.5 cM. QTL analysis of the experimental data identified seven putative genetic markers associated with hygienic behaviour. QTLs located on linkage groups 2, 4, 6 and 22 were detected for both overall hygienic behaviour and uncapping behaviour only. Individually, each QTL is of relatively small effect with each explaining only 9% � 15% of the variance in hygienic levels observed. Collectively, the putative QTLs identified here explain 79.4% of the observed variance in the expression of hygienic behaviour. These results indicate that there are many genes of low to moderate effect rather than few genes of large effect involved in this complex behavioural trait. This is typical of inherited quantitative traits which do not exhibit Mendelian phenotypic ratios. DNA extracted from the brood samples taken during testing of commercial stock, and from individual bees identified as either highly hygienic or non-hygienic in a reciprocal backcross experiment, were screened with the candidate markers associated with putative QTLs to test their diagnostic power. Unfortunately, none have produced reliably diagnostic DNA profiles. As we have now shown that hygienic behaviour is a polygenic, quantitative trait, simple diagnostic markers for Rothenbuhler's 'uncapping' and 'removal' genes are unlikely to be achieved. Our results show that the most likely way to improve disease resistance in Australian stock is via traditional methods of recurrent selection. The project was responsible for the importation of new genetic material into Australia from the United States. This hygienic stock has been well received by industry, has been widely disseminated, and incorporated into local breeding programs. We hope that it has lead to a general improvement in the level of disease resistance in Australian commercial bees.

O objetivo desse estudo foi estimar parâmetros genéticos para características produtivas e comportamentais em uma população de abelhas Apis mellifera africanizadas por meio de inferência Bayesiana. Os dados foram submetidos a análises uni e bicaracterística utilizando o programa MTGSAM. Os modelos consideraram os efeitos (fixos) de local do apiário, mês-ano ou estação-ano e o número de caixilhos com abelhas aderentes como covariável linear. As estimativas de herdabilidade apresentaram magnitudes de moderada a alta para comportamento higiênico (0,81 ± 0,17), produção de própolis (0,83 ± 0,16), produção de mel (0,37 ± 0,22) e taxa de coleta de xarope (0,39 ± 0,22) e magnitude baixa para a percentagem de ácaros em abelhas adultas (0,12 ± 0,13). A rapidez de coleta de xarope apresentou correlação genética de 0,21 ± 0,51 com produção de mel, de 0,45 ± 0,33 com produção de própolis, e de 0,05 ± 0,43 com comportamento higiênico. As correlações genéticas entre produção de mel, produção de própolis e comportamento higiênico foram de 0,20 ± 0,43, de -0,11 ± 0,41 e de 0,23 ± 0,31, respectivamente. As correlações genéticas foram negativas entre percentagem de ácaros em abelhas adultas e as características produção de mel (-0,63 ± 0,39), produção de própolis (-0,07 ± 0,50), comportamento higiênico (-0,19 ± 0,51) e rapidez de coleta de xarope (- 0,41 ± 0,51). As características produção de mel, produção de própolis e comportamento higiênico apresentam potencial para seleção genética. A menor percentagem de ácaros em abelhas adultas está relacionado a maior produção de mel e maior comportamento higiênico, mas não deve ser usado como único critério de seleção devido a baixa herdabilidade. A seleção de abelhas que coletam xarope mais rapidamente, prevendo maior produção de mel, promoverá pequeno ganho genético. Ao selecionar abelhas que produzem mais própolis haverá pequenos ganhos genéticos para comportamento higiênico ou maior produção de mel. / This study was carried out to estimate genetic parameters for productive and behavioural traits in Africanized honey bees Apis mellifera. The data were submitted uni and bicharacter analysis using the software MTGSAM. The fixed effects considered in the models were localization of the hive, month-year or season-year and number of frames covered with bees as covariate. The heritability estimates were moderate to high for hygienic behaviour (0,81 ± 0,17), propolis production (0,83 ± 0,16), honey production (0,37 ± 0,22) and syrup-collection rate (0,39 ± 0,22) and lower for percentage of mites on adult bees (0,12 ± 0,13). Syrup-collection rate showed genetic correlation values of 0,21 ± 0,51 with honey production, 0,45 ± 0,33 with propolis production and 0,05 ± 0,43 with hygienic behaviour. Genetic correlation between honey and propolis was 0,20 ± 0,43, between honey production and hygienic behaviour was -0,11 ± 0,41 and between propolis production and hygienic behaviour was 0,23 ± 0,31. Genetic correlations were negative between percentage of mites on adult bees and other traits honey production (-0,63 ± 0,39), propolis production (-0,07 ± 0,50), hygienic behaviour (-0,19 ± 0,51) and syrup-collection rate. Honey production, propolis production and hygienic behavior traits have potential for genetic selection. The lower percentage of mites on adult bees increase honey production or hygienic behaviour, but it is not recommended as the only criterion for selection, due to its low heritability. Selection for syrupcollection rate will promote small genetic gain for honey production. Propolis production is positively correlated to hygienic behaviour or honey production.

O objetivo desse estudo foi estimar parâmetros genéticos para características produtivas e comportamentais em uma população de abelhas Apis mellifera africanizadas por meio de inferência Bayesiana. Os dados foram submetidos a análises uni e bicaracterística utilizando o programa MTGSAM. Os modelos consideraram os efeitos (fixos) de local do apiário, mês-ano ou estação-ano e o número de caixilhos com abelhas aderentes como covariável linear. As estimativas de herdabilidade apresentaram magnitudes de moderada a alta para comportamento higiênico (0,81 ± 0,17), produção de própolis (0,83 ± 0,16), produção de mel (0,37 ± 0,22) e taxa de coleta de xarope (0,39 ± 0,22) e magnitude baixa para a percentagem de ácaros em abelhas adultas (0,12 ± 0,13). A rapidez de coleta de xarope apresentou correlação genética de 0,21 ± 0,51 com produção de mel, de 0,45 ± 0,33 com produção de própolis, e de 0,05 ± 0,43 com comportamento higiênico. As correlações genéticas entre produção de mel, produção de própolis e comportamento higiênico foram de 0,20 ± 0,43, de -0,11 ± 0,41 e de 0,23 ± 0,31, respectivamente. As correlações genéticas foram negativas entre percentagem de ácaros em abelhas adultas e as características produção de mel (-0,63 ± 0,39), produção de própolis (-0,07 ± 0,50), comportamento higiênico (-0,19 ± 0,51) e rapidez de coleta de xarope (- 0,41 ± 0,51). As características produção de mel, produção de própolis e comportamento higiênico apresentam potencial para seleção genética. A menor percentagem de ácaros em abelhas adultas está relacionado a maior produção de mel e maior comportamento higiênico, mas não deve ser usado como único critério de seleção devido a baixa herdabilidade. A seleção de abelhas que coletam xarope mais rapidamente, prevendo maior produção de mel, promoverá pequeno ganho genético. Ao selecionar abelhas que produzem mais própolis haverá pequenos ganhos genéticos para comportamento higiênico ou maior produção de mel. / This study was carried out to estimate genetic parameters for productive and behavioural traits in Africanized honey bees Apis mellifera. The data were submitted uni and bicharacter analysis using the software MTGSAM. The fixed effects considered in the models were localization of the hive, month-year or season-year and number of frames covered with bees as covariate. The heritability estimates were moderate to high for hygienic behaviour (0,81 ± 0,17), propolis production (0,83 ± 0,16), honey production (0,37 ± 0,22) and syrup-collection rate (0,39 ± 0,22) and lower for percentage of mites on adult bees (0,12 ± 0,13). Syrup-collection rate showed genetic correlation values of 0,21 ± 0,51 with honey production, 0,45 ± 0,33 with propolis production and 0,05 ± 0,43 with hygienic behaviour. Genetic correlation between honey and propolis was 0,20 ± 0,43, between honey production and hygienic behaviour was -0,11 ± 0,41 and between propolis production and hygienic behaviour was 0,23 ± 0,31. Genetic correlations were negative between percentage of mites on adult bees and other traits honey production (-0,63 ± 0,39), propolis production (-0,07 ± 0,50), hygienic behaviour (-0,19 ± 0,51) and syrup-collection rate. Honey production, propolis production and hygienic behavior traits have potential for genetic selection. The lower percentage of mites on adult bees increase honey production or hygienic behaviour, but it is not recommended as the only criterion for selection, due to its low heritability. Selection for syrupcollection rate will promote small genetic gain for honey production. Propolis production is positively correlated to hygienic behaviour or honey production.

O objetivo desse estudo foi estimar parâmetros genéticos para características produtivas e comportamentais em uma população de abelhas Apis mellifera africanizadas por meio de inferência Bayesiana. Os dados foram submetidos a análises uni e bicaracterística utilizando o programa MTGSAM. Os modelos consideraram os efeitos (fixos) de local do apiário, mês-ano ou estação-ano e o número de caixilhos com abelhas aderentes como covariável linear. As estimativas de herdabilidade apresentaram magnitudes de moderada a alta para comportamento higiênico (0,81 ± 0,17), produção de própolis (0,83 ± 0,16), produção de mel (0,37 ± 0,22) e taxa de coleta de xarope (0,39 ± 0,22) e magnitude baixa para a percentagem de ácaros em abelhas adultas (0,12 ± 0,13). A rapidez de coleta de xarope apresentou correlação genética de 0,21 ± 0,51 com produção de mel, de 0,45 ± 0,33 com produção de própolis, e de 0,05 ± 0,43 com comportamento higiênico. As correlações genéticas entre produção de mel, produção de própolis e comportamento higiênico foram de 0,20 ± 0,43, de -0,11 ± 0,41 e de 0,23 ± 0,31, respectivamente. As correlações genéticas foram negativas entre percentagem de ácaros em abelhas adultas e as características produção de mel (-0,63 ± 0,39), produção de própolis (-0,07 ± 0,50), comportamento higiênico (-0,19 ± 0,51) e rapidez de coleta de xarope (- 0,41 ± 0,51). As características produção de mel, produção de própolis e comportamento higiênico apresentam potencial para seleção genética. A menor percentagem de ácaros em abelhas adultas está relacionado a maior produção de mel e maior comportamento higiênico, mas não deve ser usado como único critério de seleção devido a baixa herdabilidade. A seleção de abelhas que coletam xarope mais rapidamente, prevendo maior produção de mel, promoverá pequeno ganho genético. Ao selecionar abelhas que produzem mais própolis haverá pequenos ganhos genéticos para comportamento higiênico ou maior produção de mel. / This study was carried out to estimate genetic parameters for productive and behavioural traits in Africanized honey bees Apis mellifera. The data were submitted uni and bicharacter analysis using the software MTGSAM. The fixed effects considered in the models were localization of the hive, month-year or season-year and number of frames covered with bees as covariate. The heritability estimates were moderate to high for hygienic behaviour (0,81 ± 0,17), propolis production (0,83 ± 0,16), honey production (0,37 ± 0,22) and syrup-collection rate (0,39 ± 0,22) and lower for percentage of mites on adult bees (0,12 ± 0,13). Syrup-collection rate showed genetic correlation values of 0,21 ± 0,51 with honey production, 0,45 ± 0,33 with propolis production and 0,05 ± 0,43 with hygienic behaviour. Genetic correlation between honey and propolis was 0,20 ± 0,43, between honey production and hygienic behaviour was -0,11 ± 0,41 and between propolis production and hygienic behaviour was 0,23 ± 0,31. Genetic correlations were negative between percentage of mites on adult bees and other traits honey production (-0,63 ± 0,39), propolis production (-0,07 ± 0,50), hygienic behaviour (-0,19 ± 0,51) and syrup-collection rate. Honey production, propolis production and hygienic behavior traits have potential for genetic selection. The lower percentage of mites on adult bees increase honey production or hygienic behaviour, but it is not recommended as the only criterion for selection, due to its low heritability. Selection for syrupcollection rate will promote small genetic gain for honey production. Propolis production is positively correlated to hygienic behaviour or honey production.

Ce travail porte sur la dispersion des graines par les fourmis et se divise en deux parties : l’une expérimentale (Chapitres 1, 2 & 3) et l’autre synthétique (Chapitres 4 & 5). L’approche expérimentale a consisté en une exploration en conditions de laboratoire des comportements des fourmis à deux étapes du processus de myrmécochorie: à la source de graines et dans le nid. Des graines des plantes myrmécochores Viola odorata et Chelidonium majus ont été présentées aux fourmis Lasius niger et Myrmica rubra. Chaque étape de la séquence myrmécochorique a généré une variabilité des comportements propre à chacun des quatre couples fourmis-graines. L’élaiosome n’attire pas les fourmis à distance. Les fourmis suivent toujours la même séquence de comportements : antennation, manipulation et prise des graines. Le nombre d’antennations et de manipulations avant la prise de graines peut être considéré comme un indicateur de l’« hésitation » des fourmis à prendre les graines. L’espèce à tendance carnivore Myrmica rubra a été plus rapide et efficace dans la prise de graines que l’espèce éleveuse de pucerons Lasius niger. Parallèlement, les fourmis ont moins antenné, moins manipulé et plus pris de graines de Chelidonium majus, ce qui montre un intérêt particulier pour cette espèce. Un jour après l’expérience, toutes les graines des deux espèces se trouvaient dans les déchets à l’extérieur du nid, avec au moins la moitié des élaiosomes consommés (Chapitre 1). Lors du passage des graines à l’intérieur du nid, les fourmis Myrmica rubra ont également montré une capacité à traiter les graines rapidement, en montrant une dynamique d’arrachage de l’élaiosome et de rejet des graines hors du nid plus rapide. Le taux d’arrachage de l’élaiosome a été influencé par l’espèce de graine, plus important pour les graines de Chelidonium majus. Nous avons montré qu’une proportion variable de graines rapportées au nid (moins de la moitié) étaient déposées directement au contact des larves, les autres étant traitées ailleurs par les ouvrières ou laissées temporairement à l’abandon dans le nid. Par ailleurs, les dynamiques de rejet des items hors du nid ont curieusement été peu influencées par l’espèce de graine. Pour une graine, le fait de ne plus avoir d’élaiosome diminue le nombre moyen d’ouvrières qui la contacte simultanément. Parallèlement, même si la réponse n’est pas de type « tout ou rien », l’absence d’élaiosome accroit aussi la probabilité qu’une graine a d’être rejetée. (Chapitre 2). Nous avons isolé expérimentalement le paramètre de dessiccation des graines afin mesurer son influence sur le taux de prises. La dessiccation progressive des graines réduit les taux de prises par les fourmis Myrmica rubra. La réhydratation des mêmes graines leur permet de retrouver une attractivité et donc une valeur fonctionnelle prolongée. Les graines de Viola odorata perdent presque toute attractivité après 4 jours de dessiccation et leur réhydratation ne rétabli que faiblement leur attractivité. A l’inverse, les graines de Chelidonium majus gardent un tiers de leur attractivité après un mois de dessiccation et leur réhydratation restaure presque entièrement leur attractivité (Chapitre 3). La synthèse bibliographique a permis de dresser un aperçu des principales caractéristiques des fourmis dispersatrices de graines myrmécochores. Certains traits « généralistes » rendent les rencontres entre fourmis et graines très probables : leur ubiquité et diversité taxonomique, leur régime alimentaire omnivore, et leur fourragement « diffus » et opportuniste. Les fourmis possèdent des traits qui les rendent uniques par rapport aux autres insectes : le fourragement au sol, la capacité à transporter de la nourriture, ainsi que la nidification. Certains traits des fourmis ont une influence considérable sur la dispersion des graines : leur taille, les préférences de régime alimentaire, la phénologie, la capacité d’apprentissage et la fréquence de déménagement des nids. Nous développons également l’hypothèse que la rapidité et l’efficacité du traitement des graines par les fourmis seraient une conséquence d’un comportement hygiénique des fourmis à tendance carnivore, habituées à gérer des proies périssables (Chapitre 4). Nous avons dressé pour la première fois une liste des espèces de plantes myrmécochores et potentiellement myrmécochores des régions d’Europe tempérée (260 spp.). Nous montrons que ces dernières sont majoritairement herbacées, et ont tendance à fleurir plus précocement que les autres espèces. La proportion de graines myrmécochores comprises entre 1 et 3 mm et entre 0,6 et 10 mg est plus importante que dans le reste de la flore (Chapitre 5). / This study concerns seed dispersal by ants and is divided in two parts: one experimental (chapters 1, 2 & 3), and one synthetic (chapters 4 & 5). Experimental work consisted in a series of laboratory experiments, in which ant behaviour was studied at two stages of the dispersal process: at the seed source and inside the nest. Seeds of the myrmecochorous plants Viola odorata and Chelidonium majus were presented to two ant species: Lasius niger and Myrmica rubra. Each stage of the myrmecochory sequence generated a variability of behaviours for each of the four ant-seed pairs. The elaiosomedo not attract seed at a distance. Ants followed always the same behavioural sequence: antennations, manipulations, and removal. The number of antennations and manipulations before removal can be considered as a measure of ants’ “hesitation” to remove seeds. The carnivorous species Myrmica rubra was faster and more efficient at taking seeds than the aphid-tending Lasius niger. At the same time, ants antennated and manipulated less Chelidonium majus seeds, which shows a particular interest for this seed species. One day after the experiment, all seeds of both species were located outside the nest in the refuse piles. At least half of their elaiosomes had been consumed (chapter 1). Inside the nest, Myrmica rubra also showed a great ability to treat seeds quickly, i.e. quicker dynamics of elaiosome removal and seed rejection outside the nest. Elaiosome removal rates were influenced by seed species (higher for Chelidonium majus). We showed that a variable proportion of seeds (less than half) was directly deposited in contact with larvae. The rest of the seeds were handled elsewhere by workers, or left temporarily unattended in the nest. Dynamics of seed rejection outside the nest were curiously little influenced by the seed species. For a seed, to loose the elaiosome decreased the number of workers manipulating it. At the same time, even if the rejection response is not automatic, the absence of elaiosome increases the probability for a seed to be rejected (Chapter 2). We isolated the desiccation parameter in order to measure its influence on seed removal rates. Progressive seed desiccation reduced Myrmica rubra removal rates. Rehydration of the same seeds restored their attractiveness, thereby prolonging their functional life. Viola odorata seeds lost almost all their attractiveness after 4 days of desiccation, and rehydration only restored a reduced part of their attractiveness. On the contrary, Chelidonium majus seeds kept one third of their attractiveness after one month of desiccation, and recovered almost all their attractiveness after rehydration (Chapter 3). The bibliographic review allowed us to compile an outline of the main features of seed-disperser ants. Some generalist features highly increase the probability that ants encounter seeds: their ubiquity and taxonomic diversity, omnivorous diet and their opportunistic “diffuse” foraging. Among insects, ants have unique traits that make them broad dispsersers: ground foraging, the ability to transport items, and nesting behaviour. Some other traits have a great influence on the seed dispersal system: the ant body size, their diet preferences, the phenology of the colony, the learning, and the frequence of nest relocation. We also develop the hypothesis according to which, handling efficiency of ants is a byproduct of hygienic behaviour of carnivorous oriented species, since they are used to manage perishable preys (Chapter 4). We compiled the first list of myrmecochorous and potentially myrmecochorous plants species of European temperate regions (260 spp.). We show that these plants are mainly herbaceous. They also tend to flower earlier than the whole flora. The proportions of myrmecochorous seeds having a size range between 1 and 3 mm, and a weight range between 0.6 and 10 mg are higher than in the rest of the flora (Chapter 5).

Ce travail porte sur la dispersion des graines par les fourmis et se divise en deux parties :l’une expérimentale (Chapitres 1, 2 & 3) et l’autre synthétique (Chapitres 4 & 5). L’approche expérimentale a consisté en une exploration en conditions de laboratoire des comportements des fourmis à deux étapes du processus de myrmécochorie: à la source de graines et dans le nid. Des graines des plantes myrmécochores Viola odorata et Chelidonium majus ont été présentées aux fourmis Lasius niger et Myrmica rubra. Chaque étape de la séquence myrmécochorique a généré une variabilité des comportements propre à chacun des quatre couples fourmis-graines. <p>L’élaiosome n’attire pas les fourmis à distance. Les fourmis suivent toujours la même séquence de comportements :antennation, manipulation et prise des graines. Le nombre d’antennations et de manipulations avant la prise de graines peut être considéré comme un indicateur de l’« hésitation » des fourmis à prendre les graines. L’espèce à tendance carnivore Myrmica rubra a été plus rapide et efficace dans la prise de graines que l’espèce éleveuse de pucerons Lasius niger. Parallèlement, les fourmis ont moins antenné, moins manipulé et plus pris de graines de Chelidonium majus, ce qui montre un intérêt particulier pour cette espèce. Un jour après l’expérience, toutes les graines des deux espèces se trouvaient dans les déchets à l’extérieur du nid, avec au moins la moitié des élaiosomes consommés (Chapitre 1).<p>Lors du passage des graines à l’intérieur du nid, les fourmis Myrmica rubra ont également montré une capacité à traiter les graines rapidement, en montrant une dynamique d’arrachage de l’élaiosome et de rejet des graines hors du nid plus rapide. Le taux d’arrachage de l’élaiosome a été influencé par l’espèce de graine, plus important pour les graines de Chelidonium majus. Nous avons montré qu’une proportion variable de graines rapportées au nid (moins de la moitié) étaient déposées directement au contact des larves, les autres étant traitées ailleurs par les ouvrières ou laissées temporairement à l’abandon dans le nid. Par ailleurs, les dynamiques de rejet des items hors du nid ont curieusement été peu influencées par l’espèce de graine. Pour une graine, le fait de ne plus avoir d’élaiosome diminue le nombre moyen d’ouvrières qui la contacte simultanément. Parallèlement, même si la réponse n’est pas de type « tout ou rien », l’absence d’élaiosome accroit aussi la probabilité qu’une graine a d’être rejetée. (Chapitre 2).<p>Nous avons isolé expérimentalement le paramètre de dessiccation des graines afin mesurer son influence sur le taux de prises. La dessiccation progressive des graines réduit les taux de prises par les fourmis Myrmica rubra. La réhydratation des mêmes graines leur permet de retrouver une attractivité et donc une valeur fonctionnelle prolongée. Les graines de Viola odorata perdent presque toute attractivité après 4 jours de dessiccation et leur réhydratation ne rétabli que faiblement leur attractivité. A l’inverse, les graines de Chelidonium majus gardent un tiers de leur attractivité après un mois de dessiccation et leur réhydratation restaure presque entièrement leur attractivité (Chapitre 3).<p>La synthèse bibliographique a permis de dresser un aperçu des principales caractéristiques des fourmis dispersatrices de graines myrmécochores. Certains traits « généralistes » rendent les rencontres entre fourmis et graines très probables :leur ubiquité et diversité taxonomique, leur régime alimentaire omnivore, et leur fourragement « diffus » et opportuniste. Les fourmis possèdent des traits qui les rendent uniques par rapport aux autres insectes :le fourragement au sol, la capacité à transporter de la nourriture, ainsi que la nidification. Certains traits des fourmis ont une influence considérable sur la dispersion des graines :leur taille, les préférences de régime alimentaire, la phénologie, la capacité d’apprentissage et la fréquence de déménagement des nids. Nous développons également l’hypothèse que la rapidité et l’efficacité du traitement des graines par les fourmis seraient une conséquence d’un comportement hygiénique des fourmis à tendance carnivore, habituées à gérer des proies périssables (Chapitre 4).<p>Nous avons dressé pour la première fois une liste des espèces de plantes myrmécochores et potentiellement myrmécochores des régions d’Europe tempérée (260 spp.). Nous montrons que ces dernières sont majoritairement herbacées, et ont tendance à fleurir plus précocement que les autres espèces. La proportion de graines myrmécochores comprises entre 1 et 3 mm et entre 0,6 et 10 mg est plus importante que dans le reste de la flore (Chapitre 5).<p>/<p>This study concerns seed dispersal by ants and is divided in two parts: one experimental (chapters 1, 2 & 3), and one synthetic (chapters 4 & 5). <p>Experimental work consisted in a series of laboratory experiments, in which ant behaviour was studied at two stages of the dispersal process: at the seed source and inside the nest. Seeds of the myrmecochorous plants Viola odorata and Chelidonium majus were presented to two ant species: Lasius niger and Myrmica rubra. Each stage of the myrmecochory sequence generated a variability of behaviours for each of the four ant-seed pairs.<p>The elaiosomedo not attract seed at a distance. Ants followed always the same behavioural sequence: antennations, manipulations, and removal. The number of antennations and manipulations before removal can be considered as a measure of ants’ “hesitation” to remove seeds. The carnivorous species Myrmica rubra was faster and more efficient at taking seeds than the aphid-tending Lasius niger. At the same time, ants antennated and manipulated less Chelidonium majus seeds, which shows a particular interest for this seed species. One day after the experiment, all seeds of both species were located outside the nest in the refuse piles. At least half of their elaiosomes had been consumed (chapter 1). <p>Inside the nest, Myrmica rubra also showed a great ability to treat seeds quickly, i.e. quicker dynamics of elaiosome removal and seed rejection outside the nest. Elaiosome removal rates were influenced by seed species (higher for Chelidonium majus). We showed that a variable proportion of seeds (less than half) was directly deposited in contact with larvae. The rest of the seeds were handled elsewhere by workers, or left temporarily unattended in the nest. Dynamics of seed rejection outside the nest were curiously little influenced by the seed species. For a seed, to loose the elaiosome decreased the number of workers manipulating it. At the same time, even if the rejection response is not automatic, the absence of elaiosome increases the probability for a seed to be rejected (Chapter 2).<p>We isolated the desiccation parameter in order to measure its influence on seed removal rates. Progressive seed desiccation reduced Myrmica rubra removal rates. Rehydration of the same seeds restored their attractiveness, thereby prolonging their functional life. Viola odorata seeds lost almost all their attractiveness after 4 days of desiccation, and rehydration only restored a reduced part of their attractiveness. On the contrary, Chelidonium majus seeds kept one third of their attractiveness after one month of desiccation, and recovered almost all their attractiveness after rehydration (Chapter 3). <p>The bibliographic review allowed us to compile an outline of the main features of seed-disperser ants. Some generalist features highly increase the probability that ants encounter seeds: their ubiquity and taxonomic diversity, omnivorous diet and their opportunistic “diffuse” foraging. Among insects, ants have unique traits that make them broad dispsersers: ground foraging, the ability to transport items, and nesting behaviour. Some other traits have a great influence on the seed dispersal system: the ant body size, their diet preferences, the phenology of the colony, the learning, and the frequence of nest relocation. We also develop the hypothesis according to which, handling efficiency of ants is a byproduct of hygienic behaviour of carnivorous oriented species, since they are used to manage perishable preys (Chapter 4). <p>We compiled the first list of myrmecochorous and potentially myrmecochorous plants species of European temperate regions (260 spp.). We show that these plants are mainly herbaceous. They also tend to flower earlier than the whole flora. The proportions of myrmecochorous seeds having a size range between 1 and 3 mm, and a weight range between 0.6 and 10 mg are higher than in the rest of the flora (Chapter 5). <p><p> / Doctorat en Sciences / info:eu-repo/semantics/nonPublished