Abstract Between October 1999 and October 2003, 30 adult and 48 young (< 1 year) raccoon dogs (Nyctereutes procyonoides) were monitored using radio-telemetry in an area of Germany which has been occupied by this invasive alien species since the early 1990s. Additionally, three pairs of raccoon dogs were observed by continuous radio-tracking during the first six weeks after parturition in 2003. Furthermore 136 raccoon dog pubs were ear-tagged between June 1999 and August 2006. No adult animals dispersed from the area during the study period and home ranges tended to be used for several years, probably for life. The average annual home range size, calculated using 95% fixed kernel, was 382.2 ha ± 297.4 SD for females (n = 30 seasonal home ranges) and 352.4 ha ± 313.3 SD for males (n = 32 seasonal home ranges). Paired raccoon dogs had home ranges of similar size, with pair sharing the same area all year round. Raccoon dogs occupied large core areas (85% kernel) covering 81.2% of their home ranges. The home ranges were at their smallest during the mating season. The slightly larger size of home ranges in winter suggests that, due to the temperate climate, raccoon dogs do not hibernate in Germany. Males and females formed a long-term (probably lifelong) pair bond. Same-sex neighbours ignored each other and even adjacent males/females showed neither preference nor avoidance. Thus, it can be assumed that the raccoon dog in Central Europe is monogamous without exclusive territories, based on the results of home range overlap analysis and interaction estimations. Habitat composition within home ranges and within the whole study area was almost equal. Although, percentage shares of farmland and meadow was 16.35% smaller and 12.06% higher within the home ranges, respectively. All nine habitat types (farmland, forest, settlement, water, meadows, maize fields, small woods, reeds and hedges) were used opportunistically by raccoon dogs. No significant, recognisable difference for habitat preferences between seasons was detected. Male and female raccoon dog showed equal habitat preference pattern. A comparison of active and inactive locations in different habitats found no remarkable differences. Habitat composition of individual home ranges was used to classify animals. If the percentage of forest within a home range exceeded 50% the individual was classified as a ‘forest type’ raccoon dog. If the percentage of forest habitats within a home range was less than 5%, the share of pastureland was mean 81.82% ± 16.92 SD. Consequently the individual was classified as a ‘agrarian type’ raccoon dog. Neither habitat preference nor habitat selection process differed between the two ‘types’. Habitat use and preference is discussed with relation to the ability of the raccoon dog to expand its range towards Western Europe. Males spent noticeably more time (40.5% of the time ±11.7 SD) alone with the pups than females (16.4% of the time ±8.5 SD). Females had noticeably larger 95% kernel home ranges (98.24 ha ±51.71 SD) than males (14.73 ha ±8.16 SD) and moved much longer daily distances (7,368 m ±2,015 SD) than males (4,094 m ±2,886 SD) in six weeks postpartum. The raccoon dogs being studied left the breeding den in the 6th week after the birth of the pups. In situ video observation showed that the male carried prey to the den to provide the female and the litter with food. A clear division of labour took place among parents during the period in which the pups were nursed: males guarded the litter in the den or in close vicinity of it, while the females foraged to satisfy their increased energy requirements. There were relocations of 59 (43.4%) ear-tagged young racoon dogs and mean distance from marking point was 13.5 km ±20.1 SD. Dispersal mortality rate was 69.5% among young raccoon dogs. Most animals (55.9%) were recovered nearer than 5 km from the marking point, whereas only 8.5% relocations were recorded further than 50 km from the marking point. There was no difference in the distances of relocations between sexes. Most (53.7%) relocations of ear-tagged young raccoon dogs were in August and September and, only 34.1% were recorded from October to April. Hunting (55 %) and traffic (27 %) were the major mortality factors. Radio-collared young raccoon dogs generally dispersed between July and September. The mean natal home range size (MCP 100%) with and without excursions was 502.6 ha ±66.4 SD (n = 9) and 92.1 ha ±66.4 SD (n = 17), respectively. There were no differences between sexes in the month of dispersal. The direction of travel for dispersing animals appeared to be random, with distances from 0.5 km to 91.2 km. A highly flexible dispersing behaviour is certainly one of the reasons which contribute to the high expansion success of the species.

Between October 1999 and October 2003, 30 adult and 48 young (< 1 year) raccoon dogs (Nyctereutes procyonoides) were monitored using radio-telemetry in an area of North-East Germany which has been occupied by this invasive alien species since the early 1990s. Additionally, three pairs of raccoon dogs were observed by continuous radio-tracking during the first six weeks after parturition in 2003. Furthermore 136 raccoon dog pubs were ear-tagged between June 1999 and August 2006. No adult animals dispersed from the area during the study period and home ranges tended to be used for several years, probably for life. The average annual home range size, calculated using 95% fixed kernel, was 382.2 ha ± 297.4 SD for females (n = 30 seasonal home ranges) and 352.4 ha ± 313.3 SD for males (n = 32 seasonal home ranges). Paired raccoon dogs had home ranges of similar size, with pair mates sharing the same area all year round. Raccoon dogs occupied large core areas (85% kernel) covering 81.2% of their home ranges. The home ranges were at their smallest during the mating season. The slightly larger size of home ranges in winter suggests that, due to the temperate climate, raccoon dogs do not hibernate in Germany. Males and females formed a long-term (probably lifelong) pair bond. Same-sex neighbours ignored each other and even adjacent males/females showed neither preference nor avoidance. Thus, it can be assumed that the raccoon dog in Central Europe is monogamous without exclusive territories, based on the results of home range overlap analysis and interaction estimations. Habitat composition within home ranges and within the whole study area was almost equal. Although, percentage shares of farmland and meadow was 16.35% smaller and 12.06% higher within the home ranges, respectively. All nine habitat types (farmland, forest, settlement, water, meadows, maize fields, small woods, reeds and hedges) were used opportunistically by raccoon dogs. No significant, recognisable difference for habitat preferences between seasons was detected. Male and female raccoon dog showed equal habitat preference pattern. A comparison of active and inactive locations in different habitats found no remarkable differences. Habitat composition of individual home ranges was used to classify animals. If the percentage of forest within a home range exceeded 50% the individual was classified as a ‘forest type’ raccoon dog. If the percentage of forest habitats within a home range was less than 5%, the share of pastureland was mean 81.82% ± 16.92 SD. Consequently the individual was classified as a ‘agrarian type’ raccoon dog. Neither habitat preference nor habitat selection process differed between the two ‘types’. Habitat use and preference is discussed with relation to the ability of the raccoon dog to expand its range towards Western Europe. Males spent noticeably more time (40.5% of the time ±11.7 SD) alone with the pups than females (16.4% of the time ±8.5 SD). Females had noticeably larger 95% kernel home ranges (98.24 ha ±51.71 SD) than males (14.73 ha ±8.16 SD) and moved much longer daily distances (7,368 m ±2,015 SD) than males (4,094 m ±2,886 SD) in six weeks postpartum. The raccoon dogs being studied left the breeding den in the 6th week after the birth of the pups. In situ video observation showed that the male carried prey to the den to provide the female and the litter with food. A clear division of labour took place among parents during the period in which the pups were nursed: males guarded the litter in the den or in close vicinity of it, while the females foraged to satisfy their increased energy requirements. There were relocations of 59 (43.4%) ear-tagged young raccoon dogs and mean distance from marking point was 13.5 km ±20.1 SD. Dispersal mortality rate was 69.5% among young raccoon dogs. Most animals (55.9%) were recovered nearer than 5 km from the marking point, whereas only 8.5% relocations were recorded further than 50 km from the marking point. There was no difference in the distances of relocations between sexes. Most (53.7%) relocations of ear-tagged young raccoon dogs were in August and September and, only 34.1% were recorded from October to April. Hunting (55 %) and traffic (27 %) were the major mortality factors. Radiocollared young raccoon dogs generally dispersed between July and September. The mean natal home range size (MCP 100%) with and without excursions was 502.6 ha ±66.4 SD (n = 9) and 92.1 ha ±66.4 SD (n = 17), respectively. There were no differences between sexes in the month of dispersal. The direction of travel for dispersing animals appeared to be random, with distances from 0.5 km to 91.2 km. A highly flexible dispersing behaviour is certainly one of the reasons which contribute to the high expansion success of the species.

Pentadesma butyracea Sabine est l’une des quatre espèces du genre Pentadesma endémique de l’Afrique. Elle est distribuée de la Sierra Léone au Gabon dans deux grands types d’habitats :les forêts denses humides discontinues du domaine guinéo-congolais (Haute- et Basse-Guinée) et le domaine soudanien du couloir sec du Dahomey (assimilé à une barrière à l’échange de gènes et d’espèces entre les deux blocs guinéo-congolais). Dans ce dernier, l’espèce se retrouve dans des galeries forestières et occupe une place capitale dans le développement socio-économique des communautés locales en raison des multiples biens et services que procurent ses produits (alimentation, médecine et pharmacopée traditionnelle, etc.). Cependant, des pressions d’origines multiples, telles que le ramassage des graines pour fabriquer du beurre, la fragmentation de l’habitat et sa destruction en faveur du maraîchage, les pratiques culturales inadaptées, les incendies, font peser de lourdes menaces sur l’espèce.<p>Le but de ce travail est d’acquérir les connaissances requises pour la conservation et la gestion durable des ressources génétiques de l’espèce. Trois objectifs ont été définis :(i) étudier la phylogéographie de l’espèce, (ii) étudier sa variabilité morphologique et génétique au Bénin et (iii) caractériser sa biologie de reproduction. En amont de ces travaux, nous avons développé onze marqueurs microsatellites nucléaires chez P. butyracea (chapitre 2). Ils ont été utilisés pour l’étude de la phylogéographie et la diversité génétique de P. butyracea (chapitres 3 et 5), ainsi que pour étudier la dépression de consanguinité et les paramètres de son système de reproduction (chapitre 7).<p>La caractérisation de la répartition spatiale des lignées génétiques de régions intergéniques de l’ADN chloroplastique et de l’ADN ribosomal (ITS) a détecté deux lignées génétiques allopatriques entre le Haut et le Bas-Guinéen, indiquant une forte différenciation génétique et un signal phylogéographique. L’analyse des microsatellites détecte trois pools géniques correspondant aux trois régions étudiées (Haute Guinée, Dahomey Gap et Basse Guinée). La diversité génétique est faible dans le Dahomey Gap, modérée dans le Haut-Guinéen et élevée dans le Bas-Guinéen. Ces résultats indiquent une séparation très ancienne des populations d’Afrique centrale et d’Afrique de l’ouest, alors que celles du Dahomey Gap pourraient résulter des forêts denses humides de l’Afrique de l’ouest lors de la période Holocène humide africaine. Dans ce couloir sec, les populations ont subi une forte dérive génétique, potentiellement due à des évènements de fondation. Au Bénin, deux groupes éco-morphologiques ont été détectés suivant un gradient nord-sud, contrastant avec deux pools géniques présentant une distribution est-ouest. <p>P. butyracea est une espèce auto-compatible majoritairement allogame. La corrélation de paternité est plus élevée aux niveaux intra-fruit vs. inter-fruits, et au sein d’une population de petite taille vs. de grande taille. Les principaux pollinisateurs au Bénin sont deux oiseaux (Cyanomitra verticalis, Cinnyris coccinigastrus) et trois abeilles (Apis mellifera, Meliponula togoensis, Hypotrigona sp.). La productivité totale en fruits augmente en fonction de l’âge de l’arbre et varie en fonction de l’année, atteignant un pic pour les arbres ayant un diamètre de 60-80 cm. Les graines sont récalcitrantes et ont une teneur en eau de 42.5 ± 2.9 %. <p>L’analyse des paramètres de reproduction et de diversité génétique, associés aux facteurs écogéographiques, nous a permis de proposer un échantillon de neuf populations représentatives de la diversité à l’échelle du Bénin, dans la perspective d’une conservation in situ. Le succès de celle-ci dépendra des efforts conjugués des communautés locales, de la recherche forestière et de la définition d’un cadre législatif par le politique pour la protection des habitats. La conservation ex situ est envisagée sous forme d’un verger rassemblant diverses origines, présentant l’intérêt supplémentaire de permettre d’étudier les contributions de la diversité génétique et de la plasticité phénotypique à la variation phénotypique. / Pentadesma butyracea Sabine is one of the four species of the endemic genus Pentadesma in Africa. The species is distributed from Sierra Leone to Gabon in two major types of habitats: the discontinuous and dense Guineo-Congolian rainforests (Upper and Lower Guinea) and the Sudanian domain of the dry corridor of Dahomey (considered as a barrier to the exchange of genes and species between Upper and Lower Guinea). In the latter, the species is found in gallery forests and plays a vital role in the socio-economic livelihood of local communities due to the various resources and services that provide its products (food, medicine and traditional, etc.). However, pressure from many sources including the collection of seeds to make butter, habitat fragmentation and its destruction for market gardening, inadequate agricultural practices, fires, are serious threats to the species.<p>The aim of this work was to acquire appropriate knowledge for the conservation and sustainable management of genetic resources of the species. Three objectives were defined (i) study the phylogeography of the species; (ii) evaluate its morphological and genetic variability in Benin; and (iii) characterize its reproductive biology. In a preliminary work, eleven nuclear microsatellite markers of P. butyracea were developed (Chapter 2). They were used for the study of phylogeography and genetic diversity of P. butyracea (chapters 3 and 5), and to study the inbreeding depression and parameters of its breeding system (Chapter 7).<p>The characterization of the genetic lineages and their spatial distribution using intergenic regions from chloroplast DNA and ribosomal DNA (ITS) region detected two allopatric genetic lineages between Upper and Lower Guinea, indicating a high genetic differentiation and a phylogeographic signal. Microsatellite markers allowed us to detect three genepools matching with the three studied regions (Upper Guinea, Dahomey-Gap and Lower Guinea). Genetic diversity was low in the Dahomey Gap, moderate in Upper Guinea and high in Lower Guinea. These results indicate an ancient separation of populations from Central and West Africa, while those from Dahomey Gap could originate West African rainforests (Upper Guinea) during the African humid Holocene period. In this dry corridor, populations experienced high genetic drift, possibly due to founding events. In Benin, two eco-morphological groups were detected following a north-south gradient, contrasting with two gene pools presenting an east-west distribution.<p>Pentadesma butyracea is a self-compatible, mainly allogamous species. The correlation of paternity was higher within-fruit vs. among-fruits, and in population of small size vs. large size. The main pollinators in Benin are two birds (Cyanomitra verticalis, Cinnyris coccinigastrus) and three bees (Apis mellifera, Meliponula togoensis, Hypotrigona sp.). Total productivity in fruit increases with tree age and varies yearly, reaching a peak for trees of 60-80 cm of diameter class. Seeds are recalcitrant (i.e. they cannot be conserved at low temperature), having a water content of 42.5 ± 2.9% at maturity.<p>The analysis of reproduction and genetics parameters, associated with eco-geographic factors, enabled us to select nine populations representative of the diversity in Benin, from the perspective of in situ conservation. The success of the latter will depend on combined efforts of local communities, forest research and an adequate legislative framework for the protection of habitats. Ex situ conservation is envisaged as an orchard assembling various origins, and would have the additional advantage of allowing to study the contribution of genetic diversity and phenotypic plasticity to phenotypic variation. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Abstract Between October 1999 and October 2003, 30 adult and 48 young (&amp;lt; 1 year) raccoon dogs (Nyctereutes procyonoides) were monitored using radio-telemetry in an area of Germany which has been occupied by this invasive alien species since the early 1990s. Additionally, three pairs of raccoon dogs were observed by continuous radio-tracking during the first six weeks after parturition in 2003. Furthermore 136 raccoon dog pubs were ear-tagged between June 1999 and August 2006. No adult animals dispersed from the area during the study period and home ranges tended to be used for several years, probably for life. The average annual home range size, calculated using 95% fixed kernel, was 382.2 ha ± 297.4 SD for females (n = 30 seasonal home ranges) and 352.4 ha ± 313.3 SD for males (n = 32 seasonal home ranges). Paired raccoon dogs had home ranges of similar size, with pair sharing the same area all year round. Raccoon dogs occupied large core areas (85% kernel) covering 81.2% of their home ranges. The home ranges were at their smallest during the mating season. The slightly larger size of home ranges in winter suggests that, due to the temperate climate, raccoon dogs do not hibernate in Germany. Males and females formed a long-term (probably lifelong) pair bond. Same-sex neighbours ignored each other and even adjacent males/females showed neither preference nor avoidance. Thus, it can be assumed that the raccoon dog in Central Europe is monogamous without exclusive territories, based on the results of home range overlap analysis and interaction estimations. Habitat composition within home ranges and within the whole study area was almost equal. Although, percentage shares of farmland and meadow was 16.35% smaller and 12.06% higher within the home ranges, respectively. All nine habitat types (farmland, forest, settlement, water, meadows, maize fields, small woods, reeds and hedges) were used opportunistically by raccoon dogs. No significant, recognisable difference for habitat preferences between seasons was detected. Male and female raccoon dog showed equal habitat preference pattern. A comparison of active and inactive locations in different habitats found no remarkable differences. Habitat composition of individual home ranges was used to classify animals. If the percentage of forest within a home range exceeded 50% the individual was classified as a ‘forest type’ raccoon dog. If the percentage of forest habitats within a home range was less than 5%, the share of pastureland was mean 81.82% ± 16.92 SD. Consequently the individual was classified as a ‘agrarian type’ raccoon dog. Neither habitat preference nor habitat selection process differed between the two ‘types’. Habitat use and preference is discussed with relation to the ability of the raccoon dog to expand its range towards Western Europe. Males spent noticeably more time (40.5% of the time ±11.7 SD) alone with the pups than females (16.4% of the time ±8.5 SD). Females had noticeably larger 95% kernel home ranges (98.24 ha ±51.71 SD) than males (14.73 ha ±8.16 SD) and moved much longer daily distances (7,368 m ±2,015 SD) than males (4,094 m ±2,886 SD) in six weeks postpartum. The raccoon dogs being studied left the breeding den in the 6th week after the birth of the pups. In situ video observation showed that the male carried prey to the den to provide the female and the litter with food. A clear division of labour took place among parents during the period in which the pups were nursed: males guarded the litter in the den or in close vicinity of it, while the females foraged to satisfy their increased energy requirements. There were relocations of 59 (43.4%) ear-tagged young racoon dogs and mean distance from marking point was 13.5 km ±20.1 SD. Dispersal mortality rate was 69.5% among young raccoon dogs. Most animals (55.9%) were recovered nearer than 5 km from the marking point, whereas only 8.5% relocations were recorded further than 50 km from the marking point. There was no difference in the distances of relocations between sexes. Most (53.7%) relocations of ear-tagged young raccoon dogs were in August and September and, only 34.1% were recorded from October to April. Hunting (55 %) and traffic (27 %) were the major mortality factors. Radio-collared young raccoon dogs generally dispersed between July and September. The mean natal home range size (MCP 100%) with and without excursions was 502.6 ha ±66.4 SD (n = 9) and 92.1 ha ±66.4 SD (n = 17), respectively. There were no differences between sexes in the month of dispersal. The direction of travel for dispersing animals appeared to be random, with distances from 0.5 km to 91.2 km. A highly flexible dispersing behaviour is certainly one of the reasons which contribute to the high expansion success of the species.

Between October 1999 and October 2003, 30 adult and 48 young (< 1 year) raccoon dogs (Nyctereutes procyonoides) were monitored using radio-telemetry in an area of North-East Germany which has been occupied by this invasive alien species since the early 1990s. Additionally, three pairs of raccoon dogs were observed by continuous radio-tracking during the first six weeks after parturition in 2003. Furthermore 136 raccoon dog pubs were ear-tagged between June 1999 and August 2006. No adult animals dispersed from the area during the study period and home ranges tended to be used for several years, probably for life. The average annual home range size, calculated using 95% fixed kernel, was 382.2 ha ± 297.4 SD for females (n = 30 seasonal home ranges) and 352.4 ha ± 313.3 SD for males (n = 32 seasonal home ranges). Paired raccoon dogs had home ranges of similar size, with pair mates sharing the same area all year round. Raccoon dogs occupied large core areas (85% kernel) covering 81.2% of their home ranges. The home ranges were at their smallest during the mating season. The slightly larger size of home ranges in winter suggests that, due to the temperate climate, raccoon dogs do not hibernate in Germany. Males and females formed a long-term (probably lifelong) pair bond. Same-sex neighbours ignored each other and even adjacent males/females showed neither preference nor avoidance. Thus, it can be assumed that the raccoon dog in Central Europe is monogamous without exclusive territories, based on the results of home range overlap analysis and interaction estimations. Habitat composition within home ranges and within the whole study area was almost equal. Although, percentage shares of farmland and meadow was 16.35% smaller and 12.06% higher within the home ranges, respectively. All nine habitat types (farmland, forest, settlement, water, meadows, maize fields, small woods, reeds and hedges) were used opportunistically by raccoon dogs. No significant, recognisable difference for habitat preferences between seasons was detected. Male and female raccoon dog showed equal habitat preference pattern. A comparison of active and inactive locations in different habitats found no remarkable differences. Habitat composition of individual home ranges was used to classify animals. If the percentage of forest within a home range exceeded 50% the individual was classified as a ‘forest type’ raccoon dog. If the percentage of forest habitats within a home range was less than 5%, the share of pastureland was mean 81.82% ± 16.92 SD. Consequently the individual was classified as a ‘agrarian type’ raccoon dog. Neither habitat preference nor habitat selection process differed between the two ‘types’. Habitat use and preference is discussed with relation to the ability of the raccoon dog to expand its range towards Western Europe. Males spent noticeably more time (40.5% of the time ±11.7 SD) alone with the pups than females (16.4% of the time ±8.5 SD). Females had noticeably larger 95% kernel home ranges (98.24 ha ±51.71 SD) than males (14.73 ha ±8.16 SD) and moved much longer daily distances (7,368 m ±2,015 SD) than males (4,094 m ±2,886 SD) in six weeks postpartum. The raccoon dogs being studied left the breeding den in the 6th week after the birth of the pups. In situ video observation showed that the male carried prey to the den to provide the female and the litter with food. A clear division of labour took place among parents during the period in which the pups were nursed: males guarded the litter in the den or in close vicinity of it, while the females foraged to satisfy their increased energy requirements. There were relocations of 59 (43.4%) ear-tagged young raccoon dogs and mean distance from marking point was 13.5 km ±20.1 SD. Dispersal mortality rate was 69.5% among young raccoon dogs. Most animals (55.9%) were recovered nearer than 5 km from the marking point, whereas only 8.5% relocations were recorded further than 50 km from the marking point. There was no difference in the distances of relocations between sexes. Most (53.7%) relocations of ear-tagged young raccoon dogs were in August and September and, only 34.1% were recorded from October to April. Hunting (55 %) and traffic (27 %) were the major mortality factors. Radiocollared young raccoon dogs generally dispersed between July and September. The mean natal home range size (MCP 100%) with and without excursions was 502.6 ha ±66.4 SD (n = 9) and 92.1 ha ±66.4 SD (n = 17), respectively. There were no differences between sexes in the month of dispersal. The direction of travel for dispersing animals appeared to be random, with distances from 0.5 km to 91.2 km. A highly flexible dispersing behaviour is certainly one of the reasons which contribute to the high expansion success of the species.