Document Type



Doctor of Philosophy (PhD)



First Advisor's Name

Hong Liu

First Advisor's Committee Title

Committee chair

Second Advisor's Name

Joel Heinen

Second Advisor's Committee Title

Committee member

Third Advisor's Name

Suzanne Koptur

Third Advisor's Committee Title

Committee member

Fourth Advisor's Name

Javier Francisco-Ortega

Fourth Advisor's Committee Title

Committee member

Fifth Advisor's Name

Scott Zona

Fifth Advisor's Committee Title

Committee member


Flora of Malesiana, Wallacea, hornbills, Moluccan Islands, dispersal syndromes, tropical trees, plant-animal interactions

Date of Defense



The dispersal services of frugivores affect plant community assembly, persistence, and gene flow in the short-term, and in the long-term are critical to ensuring that tropical trees and palms can regenerate in disturbed areas and can migrate amidst climate change. Halmahera is the largest Moluccan island within the Wallacea biodiversity hotspot, yet data on its plant and animal distributions and interactions are almost null. I studied the tropical trees and palms of Halmahera and their seed dispersal dynamics. Chapter I explores the palms of the Moluccan islands through field-, herbarium-, and literature- based studies. The results of herbarium specimen collections are presented within a preliminary list of palms for Halmahera’s Aketajawe-Lolobata National Park, and contextualized in a review of regional palm biogeography. Expanding beyond the study of one plant family, Chapter II compiles and examines all the tree and palm taxa of the Moluccan islands in order to infer seed dispersal syndromes for each taxon, resulting in an analysis of over 900 taxa. Zoochory was found in 93% of plant families, and nearly 30% of endemic taxa rely primarily on dispersal by large-bodied frugivores. The role of a hypothesized keystone disperser (the Papuan hornbill, Rhyticeros plicatus ruficollis) is confirmed experimentally to disperse about 10% of Halmahera’s tropical tree and palm taxa. The final chapter encompasses an ex-situ germination trial and a year-long in-situ experimental study that examines the germination and recruitment of three palm species post-ingestion by Papuan hornbills, in four different habitat types representative of Halmahera’s landscape. Palms dispersed by hornbills into disturbed habitats and primary forests resulted in enhanced recruitment, although results varied by habitat and species. Hornbills aid both in forest conservation and recovery/assembly after disturbance. The results of this dissertation provide a foundation for further ecological studies and for enhanced conservation of Halmahera island.





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