Escalation (macroevolutionary increase) or divergence (disparity between relatives) in trait values are two frequent outcomes of the plant-herbivore arms race. We studied the defences and caterpillars associated with 21 sympatric New Guinean figs. Herbivore generalists were concentrated on hosts with low protease and oxidative activity. The distribution of specialists correlated with phylogeny, protease and trichomes. Additionally, highly specialised Asota moths used alkaloid rich plants. The evolution of proteases was conserved, alkaloid diversity has escalated across the studied species, oxidative activity has escalated within one clade, and trichomes have diverged across the phylogeny. Herbivore specificity correlated with their response to host defences: escalating traits largely affected generalists and divergent traits specialists; but the effect of escalating traits on extreme specialists was positive. In turn, the evolution of defences in Ficus can be driven towards both escalation and divergence in individual traits, in combination providing protection against a broad spectrum of herbivores.

Bell Museum and Department of Plant and Microbial Biology University of Minnesota 250 Biological Science Center 1445 Gortner Avenue Saint Paul 55108 MN USA

Biology Centre The Czech Academy of Sciences Branisovska 31 37005 Ceske Budejovice Czech Republic

Centre for Natural Resources Research and Development University of Goroka Goroka P O Box 1078 Eastern Highland Province Papua New Guinea

Department of Biology Lund University Sölvegatan 37 223 62 Lund Sweden

Faculty of Science University of South Bohemia in Ceske Budejovice Branisovska 31 37005 Ceske Budejovice Czech Republic

Institute of Botany Czech Academy of Sciences Dukelska 135 Trebon 37982 Czech Republic

Maestria de Entomologia Universidad de Panama 080814 Panama City Panama

National Museum of Natural History Smithsonian Institution 10th St and Constitution Ave NW Washington 20560 DC USA

Natural Chemistry Research Group Department of Chemistry University of Turku FI 20014 Turku Finland

New Guinea Binatang Research Center P O Box 604 Madang Papua New Guinea

Smithsonian Tropical Research Institute Apartado 0843 03092 Balboa Ancon Panamá

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