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  • 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


    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


    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



    urrently, an estimated number of 1.5 to 30 million species live on earth (Erwin 1983; Erwin 1991; May 1988). These species are distinctive in their mor-

    phology, physiology, and behavior and coexist in regional to local communities. Community com- position is influenced by a range of ecological factors (dispersal ability and habitat selection of a species, interspecific interactions), evolutionary processes and historical events (Fig. 1).

    Differences in ecological characteristics of species lead to differences in their functionality and their role in ecosystem processes (Darwin and Wallace 1858; Loreau et al. 2001). The ques- tions of which factors structure local communities and how regional species pools contribute to local communities continue to be among the central topics in ecology (Diamond 1975).

    Following Fauth et al. (1996), animal “communities” are comprised of at least two species that interact at a given geographical location. Species “assemblages” are defined as a subset of these communities consisting of animals from similar taxa, while a species “ensemble” refers to a set of species that are closely related and use similar resources.


    Species have basic ecological requirements that have to be fulfilled to ensure their persistence in a certain local environment. Those require- ments determine - in conjunction with interspeci- fic interactions - the distribution of a species within its geographic range. The multidimen- sional combination of abiotic and biotic factors required for the persistence of a species has been considered as the “ecological niche” (Grinnell 1917; Hutchinson 1957). In contrast, MacArthur and Levins (1967) stressed the impacts of a species on its environment (e.g., its functional role within the food chain, see Elton 1927). Recent concepts combine both aspects in a single definition of the niche of a species as “the environmental conditions that allow a species to satisfy its minimum requirements so that the birth rate of a local population is equal or greater than its death rate along with the set of per capita impacts of that species on these environmental conditions” (Chase and Leibold 2003).

    Starting from the early studies of Hutchinson (1959) and MacArthur (1958; MacArthur 1972a), research efforts in community ecology have at- tempted to reveal the mechanisms that allow the coexistence of species