Structuring of animal communities: Interspecific interactions and habitat selection among ants and small mammals Habilitationsschrift zur Erlangung der Venia Legendi an der Universität Ulm Fakultät für Naturwissenschaften vorgelegt von Dr. Martin Pfeiffer Ulm, Oktober 2007 Year’s end — Still in straw hat And sandals. Basho (1644 -1694) In dieser Arbeit werden die Untersuchungen zur Gemeinschaftsökologie von Ameisen und Kleinsäugern vorgestellt, die ich zwischen 1997 und 2007 durchgeführt oder betreut habe. Ich versichere, dass ich die vorliegende Arbeit ohne fremde Hilfe angefertigt und mich keiner anderen als der ausdrücklich angegebenen Hilfsmittel bedient habe. Martin Pfeiffer Ulm, 30. Oktober 2007 CONTENTS Acknowledgments 1 1. Disentangling life histories, organization, and functions in animal communities of tropical rainforests and arid areas – an overview 3 2. Internet-based ant taxonomy and biodiversity informatics 8 3. Ant diversity gradients and faunistic inventory 14 4. Null model studies of interspecific interactions: community structure of Malaysian ants 18 5. The Sarawak soil ant project: Niches, trophic levels, and community patterns in rainforest ants 21 6. Ant- plant mutualism: Myrmecochory - seed dispersal by ants 24 7. Spatial organization in Bornean small mammal assemblages 27 8. Rainforest logging in Borneo: impacts on non-volant small mammal assemblages 30 References 34 Research articles ordered Research articles belonging to Chapter 3 43 Pfeiffer M, Chimedregzen L, Ulykpan K (2003) Community organization and species richness of ants (Hymenoptera/Formicidae) in Mongolia along an ecological gradient from steppe to Gobi desert. Journal of Biogeography 30:1921-1935 Pfeiffer M, Schultz R, Radchenko A, Yamane S, Woyciechowski M, Ulykpan A, Seifert B (2006) A critical checklist of the ants of Mongolia (Hymenoptera : Formicidae). Bonner Zoologische Beiträge 55:1-8 Paknia O, Radchenko AG, Helen A, Pfeiffer M (submitted) A Preliminary Checklist of the Ants (Hymenoptera: Formicidae) of Iran. Myrmecological News Research articles belonging to Chapter 4 81 Pfeiffer M, Ho CT, Teh CL (in press) Exploring arboreal ant community composition and co-occurrence patterns in plantations of oil palm (Elaeis guineensis) in Borneo and Peninsular Malaysia Ecography Pfeiffer M, Nais J, Linsenmair KE (2006) Worker size and seed size selection in 'seed'-collecting ant ensembles (Hymenoptera : Formicidae) in primary rain forests on Borneo. Journal of Tropical Ecology 22:685-693 Contents Research articles belonging to Chapter 6 111 Pfeiffer M, Huttenlocher H, Ayasse M (manuscript) Myrmecochory in spring ephemerals with different types of elaiosomes: how chemical composition of diaspores influences ant removal rates. Pfeiffer M, Nais J, Linsenmair KE (2004) Myrmecochory in the Zingiberaceae: seed removal of Globba franciscii and G. propinqua by ants (Hymenoptera -Formicidae) in rain forests on Borneo. Journal of Tropical Ecology 20:705-708 Research articles belonging to Chapter 7 139 Wells K, Pfeiffer M, Lakim MB, Linsenmair KE (2004) Use of arboreal and terrestrial space by a small mammal community in a tropical rainforest in Borneo, Malaysia. Journal of Biogeography 31:641-652 Wells K, Pfeiffer M, Lakim MB, Linsenmair KE (2004) Arboreal spacing patterns of the large pencil-tailed tree mouse, Chiropodomys major (Muridae), in a rainforest in Sabah, Malaysia. Ecotropica 10:15-22 Wells K, Lakim MB, Pfeiffer M (2006) Nest sites of rodents and treeshrews in Borneo. Ecotropica 12:141-149 Research articles belonging to Chapter 8 171 Wells K, Kalko EKV, Lakim MB, Pfeiffer M (2007) Effects of rain forest logging on species richness and assemblage composition of small mammals in Southeast Asia. Journal of Biogeography 34:1087-1099 Wells K, Pfeiffer M, Lakim MB, Kalko EKV (2006) Movement trajectories and habitat partitioning of small mammals in logged and unlogged rain forests on Borneo. Journal of Animal Ecology 75:1212-1223 Wells K, Kalko EKV, Lakim MB, Pfeiffer M (accepted) Movement and ranging patterns of a tropical rat in logged and unlogged rainforests Journal of Mammalogy Wells K, Smales LR, Kalko EKV, Pfeiffer M (2007) Impact of rain-forest logging on helminth assemblages in small mammals (Muridae, Tupaiidae) from Borneo. Journal of Tropical Ecology 23:35-43 Curriculum Vitae and list of publications 225 1 ACKNOWLEDGMENTS I am grateful to Prof. Dr Elisabeth Kalko for the opportunity to work as a biologist at the University of Ulm, for our productive cooperation, her constant support of my work, and for the pleasure and sense of achievement while teaching and researching at the Institute for Experimental Ecology. I also thank Prof. Dr Manfred Ayasse for our excellent cooperation and his help during the Habilitation process. I owe a debt of gratitude to Dr Marco Tschapka and Dr Konstans Wells for our great teamwork, for the many discussions about our work, and for their comments on earlier versions of this Habilitation thesis. Likewise, I thank all the members of the Institute for Experimental Ecology at the University of Ulm for their support and the inspiring atmosphere during our work, among others Dr Heiko Bellmann, Jakob Fahr, Dr Joanna Fietz, Dr Stefan Jarau, Prof. Dr Gerhard Maier, Dr Johannes Stökl and -last but not least- Ingrid Dillon. I thank in particular the members of my working group for their help and for all those fun times we had during the past six years: Katrin Deufel, Dirk Mezger, Omid Paknia, Heiko Huttenlocher and Hans Peter Katzmann. I also thank the technical staff for their various form of support: Malte Petershagen, Jolanta Schenk, Ulrike Stehle, Andrea Weiss, and Gabriele Wiest and all those who helped me during my expeditions and work in Mongolia, Malaysia, and Iran. Many thanks especially to the many students who worked in our projects and to all our collaborators and co-workers! Moreover, I am grateful to DAAD, DFG, the Gewinner-Stiftung, and the Pfeiffer family for their various financial supports of my studies. Most thankful I am to my wife Irmi, who shares my life and work in various countries, “Ohne Dich geht nix!”. This thesis has greatly benefited from the language corrections by Brian Hillcoat, Neuss. 3 DISENTANGLING LIFE HISTORIES, ORGANIZATION, AND FUNCTIONS IN ANIMAL COMMUNITIES OF TROPICAL RAINFORESTS AND ARID AREAS – AN OVERVIEW urrently, an estimated number of 1.5 to Starting from the early studies of Hutchinson 30 million species live on earth (Erwin (1959) and MacArthur (1958; MacArthur 1972a), C 1983; Erwin 1991; May 1988). These research efforts in community ecology have at- species are distinctive in their mor- tempted to reveal the mechanisms that allow the phology, physiology, and behavior and coexist in coexistence of species in local habitats and eco- regional to local communities. Community com- systems (Brown 1995). Much of the last decades position is influenced by a range of ecological of community ecology have been devoted to the factors (dispersal ability and habitat selection of a analysis and interpretation of the structure and species, interspecific interactions), evolutionary composition of animal and plant communities. processes and historical events (Fig. 1). Differences in ecological characteristics of species lead to differences in their functionality Evolutionary Processes Historical Events and their role in ecosystem processes (Darwin and Wallace 1858; Loreau et al. 2001). The ques- Regional Species Pool tions of which factors structure local communities and how regional species pools contribute to local communities continue to be among the central Habitat- Interspecific Dispersal topics in ecology (Diamond 1975). Selection Interactions Ability Following Fauth et al. (1996), animal “communities” are comprised of at least two Local species that interact at a given geographical Community Composition location. Species “assemblages” are defined as a subset of these communities consisting of animals from similar taxa, while a species “ensemble” Fig. 1 The recruitment of a local community from a regional species pool is a consequence of many inter- refers to a set of species that are closely related acting factors (Adapted from Morin 1999). and use similar resources. There are contrasting doctrines on this matter: THE NICHE CONCEPT AND ITS LIMITA- the “deterministic” view argues that competition TIONS: A SCIENTIFIC DEBATE is the main driving force in the structuring of communities. This view is linked with the “com- Species have basic ecological requirements petitive exclusion principle” as proposed by that have to be fulfilled to ensure their persistence Gause (1936) and formulated by Hardin (1960). in a certain local environment. Those require- The concept states that two species must differ in ments determine - in conjunction with interspeci- some traits related to their fitness in order to co- fic interactions - the distribution of a species exist. Consequently, coexisting organisms should within its geographic range. The multidimen- differ in their ecological requirements (Biesmeijer sional combination of abiotic and biotic factors et al. 1999; Gotelli and McCabe 2002; Graves required for the persistence of a species has been and Gotelli 1993). As part of this view, considered as the “ecological niche” (Grinnell community structure is seen as a non-random 1917; Hutchinson 1957). In contrast, MacArthur association of locally coexisting species (Brown and Levins (1967) stressed the impacts of a 1995). Furthermore, assembly rules have been species on its environment (e.g.,