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Seed Dispersal and Plant–Gorilla Mutualism in Southeast Cameroon

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Seed Dispersal and Plant–Gorilla Mutualism in Southeast Cameroon COMMUNAUTÉ FRANÇAISE DE BELGIQUE UNIVERSITÉ DE LIÈGE – GEMBLOUX AGRO-BIO TECH EFFECTIVENESS OF WESTERN LOWLAND GORILLA (GORILLA GORILLA GORILLA) SEED DISPERSAL AND PLANT–GORILLA MUTUALISM IN SOUTHEAST CAMEROON Charles-Albert PETRE Dissertation originale présentée en vue de l’obtention du grade de docteur en sciences agronomiques et ingénierie biologique Promoteurs : Professeur Jean-Louis Doucet & Docteur Roseline Claire Beudels-Jamar Année 2016 Institut Royal des Sciences Naturelles de Belgique Société Royale de Zoologie d’Anvers COMMUNAUTÉ FRANÇAISE DE BELGIQUE UNIVERSITÉ DE LIÈGE – GEMBLOUX AGRO-BIO TECH EFFECTIVENESS OF WESTERN LOWLAND GORILLA (GORILLA GORILLA GORILLA) SEED DISPERSAL AND PLANT–GORILLA MUTUALISM IN SOUTHEAST CAMEROON Charles-Albert PETRE Dissertation originale présentée en vue de l’obtention du grade de docteur en sciences agronomiques et ingénierie biologique Promoteurs : Professeur Jean-Louis Doucet & Docteur Roseline Claire Beudels-Jamar Année 2016 Institut Royal des Sciences Naturelles de Belgique Société Royale de Zoologie d’Anvers Copyright. Aux termes de la loi belge du 30 juin 1994 sur le droit d’auteur et les droits voisins, seul l’auteur a le droit de reproduire partiellement ou complètement cet ouvrage de quelque façon et forme que ce soit ou d’en autoriser la reproduction partielle ou complète de quelque manière et sous quelque forme que ce soit. Toute photocopie ou reproduction sous autre forme est donc faite en violation de ladite loi et des modifications ultérieures. Copyright. According to the Belgian law of the 30th of June 1994, on the author right and associated rights, only the author has the right to partially or fully reproduce this manuscript in any way and any form possible and to authorisze the partial or full reproduction in any way and any form possible. Any copy or reproduction in a different way and form than above is therefore against the law and any modifications of this law. Petre, Charles-Albert (2016). Efficacité de la dispersion des graines par le gorille des plaines de l’ouest (Gorilla gorilla gorilla) et relations plante–gorille mutualistes au sud-est Cameroun. Thèse de doctorat. Université de Liège–Gembloux Agro-Bio-Tech, 155p. Résumé Les frugivores de grande taille jouent un rôle fondamental dans la dynamique forestière et occupent une place déterminante dans les réseaux de dispersion de graines. Cependant, la grande faune est davantage vulnérable aux extinctions locales résultant des perturbations anthropiques grandissantes. Les changements observés dans la structure et la composition du jeune recrû après la disparition des grands frugivores suggèrent une modification sur le long terme de l'écosystème forestier qui pourrait se traduire par une perte de biodiversité et une réduction des services écosystémiques. Le gorille des plaines de l’ouest (Gorilla gorilla gorilla Savage, 1847) est le plus grand des frugivores arboricoles, et il subit malheureusement des réductions populationnelles localement dramatiques. Afin de mieux comprendre les conséquences écologiques de leur disparition, la présente thèse s’attelle à identifier et décrire les différentes composantes de l’efficacité de la dispersion des graines chez une population de gorilles des plaines de l’ouest (site "La Belgique", dans le sud-est du Cameroun). Cinquante-huit espèces ont été retrouvées dans 1030 fèces collectées pendant 36 mois. Malgré cette diversité, les espèces du genre Uapaca (N=5), dont les graines étaient présentes dans les fèces durant 72.2% des mois échantillonnés, contribuaient à elles seules à 36.5% de la quantité totale de graines dispersées durant la période d’étude. Avec un temps moyen de rétention des graines dans le tractus digestif de 50.6 ± 28.1 h et une distance moyenne parcourue quotidiennement de 2 km, les distances de dispersion sont potentiellement très élevées. Cette contribution à la dynamique de métapopulation serait d’autant plus effective que le pouvoir germinatif des graines ne se réduit pas avec le temps de rétention (N=4 spp.), et peut même en être amélioré (N=1 sp.). De façon générale, les graines reçoivent un bon traitement dans le tractus digestif puisque 50% et 41.7% des espèces testées (N=12) possèdaient un pouvoir germinatif après transit inchangé (dont Uapaca spp.) et accru, respectivement. Au niveau des sites de déposition des graines, les bousiers (Scarabaeidae coprophages) peuvent toutefois l'altérer en enterrant trop profondément les graines, comme démontré avec une espèce du genre Uapaca (73% des graines enfouies se retrouvaient à >10 cm de profondeur où la probabilité d’émergence était nulle). Cet effet négatif des bousiers est cependant moindre pour les graines dispersées dans les sites de nidification, où l’activité des bousiers moins importante résulte en une plus faible proportion de graines enfouies et à de plus faibles profondeurs (8.7 ± 3.9 cm). Les sites de nidification constituent des sites de déposition importants puisque la moitié des fèces y sont produites, et sont caractérisés par une végétation arborée à canopée basse et éparse (ouverture moyenne de 19.1 ± 13.1%) et dont le tempérament héliophile et la classe de diamètre 10–19.9 cm sont surreprésentés, témoignant de perturbations récentes telles que la i chute d’arbres émergents. Les conditions lumineuses favorables rencontrées dans ces microhabitats favorisent le recrutement des espèces végétales dispersées, comme démontré avec les espèces du genre Uapaca. Les probabilités de recrutement de ces dernières sont davantage accrues lorsque les sites de nidification sont établis sur sol hygromorphe, habitat prépondéramment utilisé pour la nidification en fin de grande saison sèche (50% des sites de nidification rencontrés à cette période étaient construits dans ce type d’habitat). Puisque cette période correspond au pic de dispersion des graines du genre Uapaca (moyenne de 31.7 ± 25.5 graines par 100 g de matière fécale ; 38.6% des graines ont été dispersées à cette période), la population de gorilles du site La Belgique réalisent une ‘dispersion dirigée’, faisant de ce disperseur un contributeur particulièrement efficace de la dynamique de population de ce genre. En contrepartie, les fruits du genre Uapaca constituent une ressource alimentaire, dites de soudure, contribuant au fitness des gorilles d’une façon non égalée par les fruits des autres espèces, mettant en évidence un cas de mutualisme plante–animale. Cette thèse confirme donc le rôle capital des larges frugivores dans le façonnement des forêts tropicales et le maintien de leur intégrité. Des actions appropriées de gestion des forêts tropicales doivent être urgemment entreprises pour éviter que ces mammifères clés ne disparaissent. Mots clés : comportement nidificateur, dispersion dirigée, dispersion secondaire, dynamique forestière, endozoochorie, frugivores de grandes tailles, recrutement de jeunes plants, site de nidification, Uapaca. ii Petre, Charles-Albert (2016). Efficacité de la dispersion des graines par le gorille des plaines de l’ouest (Gorilla gorilla gorilla) et relations plante–gorille mutualistes au sud-est Cameroun. Thèse de doctorat. Université de Liège–Gembloux Agro-Bio-Tech, 155p. Summary Large frugivores contribute a notable role in forest dynamics and hence occupy a central position in seed dispersal networks. However, large-bodied vertebrates are disproportionately prone to local extinction caused by increasing human disturbances than are smaller-bodied species. Observed changes in the structure and composition of juvenile plant cohorts following the extirpation of large-bodied seed dispersers constitute evidence for the long-term alteration of the whole tree stand, resulting in a loss of biodiversity and a reduction of ecosystem services. The western lowland gorilla (Gorilla gorilla gorilla Savage, 1847) is the largest arboreal frugivore, facing rapid population depletion in recent years and becoming locally extinct in many parts of its historical range. To understand the likely consequence of their disappearance for the future of Afrotropical forests, this thesis aims at determining the seed dispersal effectiveness of a western lowland gorilla population (“La Belgique” research site, southeast Cameroon). Seeds of 58 species were identified in 1030 faecal samples collected over 36 months. Despite this diversity of species dispersed, seeds of the genus Uapaca (N=5), dispersed in 72.2% of months sampled, represented 36.5% of the total seed load. With a retention time of seeds in the gorilla digestive tract averaging 50.6 ± 28.1 h, and an average daily travel distance of 2 km, the western lowland gorilla may transport seeds over considerably long distances. This contribution to (meta)population dynamics would be effective since seed viability is not reduced by the time seeds are exposed to digestive enzymes (N=4 spp.) and may even be improved (N=1 sp.). Overall, seeds received a good treatment in the mouth and gut as 50% and 41.7% of species tested (N=12 spp.) had a germination success unchanged (including Uapaca spp.) or improved, respectively, after gut passage. At the seed deposition site, however, seed viability may be further altered by dung beetles (coprophageous Scarabaeidae) through burial at unsuitable depths, as demonstrated by one species of Uapaca (73% of buried seeds were relocated >10 cm deep where the emergence probability dropped to zero). This negative impact is nonetheless lower at sleeping sites where fewer seeds are buried and at shallower depths (mean depth: 8.7 ± 3.9 cm). Sleeping sites are important
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