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Summary: Survival benefits select for group living in a social spider despite
reproductive costs
T. BILDE,*,§ K. S. COATES,* K. BIRKHOFER, T. BIRD,ŕ A. A. MAKLAKOV,§, Y. LUBIN** &
L. AVILE´ S
*Department of Biological Sciences, Ecology and Genetics, University of Aarhus, Aarhus, Denmark
Institut fu¨r Zoologie, TU Darmstadt Schnittspahnstr, Darmstadt, Germany
ŕNational Museum of Namibia, Windhoek, Namibia
§Department of Ecology and Evolution, Evolutionary Biology Centre, Uppsala University, Uppsala, Sweden
Department of Life Sciences, Ben Gurion University, Beer Sheva, Israel
**Mitrani Department of Desert Ecology, Blaustein Institute for Desert Research, Ben-Gurion University, Sede Boker, Israel
Department of Zoology, University of British Columbia, Vancouver, BC, Canada
Introduction
The evolution of cooperation in animals raises the
puzzling problem of how helping is maintained despite
the inherent costs associated with it. Solving this prob-
lem, which has proven one of the greatest challenges in
modern biology, is central to understanding the selective
forces driving the association of lower level entities into
higher levels of organization (Maynard Smith & Szathm-
a´ry, 1995; Reeve & Keller, 1999). It is generally assumed
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