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The most conspicuous and species-rich ant-plant mutualism in the Malesian region is found in the important pioneer tree genus Macurungu, yet little is known about the identities or community ecology of the species involved. Our studies have revealed a far more complex system than previously thought. This paper presents the first extensive investigation in the whole distribution area of myrmecophytic Mucarungu. All ant-inhabited species were restricted to the moister parts of SE Asia: Peninsular Malaysia, South and East Thailand, Sumatra and Borneo. We found a rather strict and similar altitudinal zonation of myrmecophytic Macaranga species in all regions. Here we focus on the majority of the 19 Mucaranga species obligatorily associated with ants of the genus Crematogater. We identified a total of 2163 ant queens which belonged to at least eight (morpho)species of the small subgenus Decacraa as well as to one non-Decacrau @robably from Atopogyne). The ant species were not randomly distributed among the Mucuranga species but distinct patterns of associations emerged. Despite common sympatric distribution of Mucurangu species, in most cases a surprisingly high specificity of ant colonization was maintained which was, however, often not species-specific but groups of certain plant species with identical ant partners could be found. These colonization patterns usually but not always mirror existing taxonomic sections within the genus Macaranga. Possible mechanisms of specificity are discussed. The results are compared with other ant-plant mutualisms. 0 1999 The Linnran Socirtv of Ixlndon ADDITIONAL KEY WORDS:-biogeography ~ co-evolution -Decucremuhost specificity~

Summary. In the humid tropics of SE Asia there are some 14 myrmecophytic species of the pioneer tree genus M acaranga (Euphorbiaceae). In Peninsular Malaysia a close association exists between the trees and the small, non-stinging myrmicine Crema togas ter borneensis. These ants feed mainly on food bodies provided by the plants and have their colonies inside the hollow intemodes. In a ten months field study we were able to demonstrate for four Macaranga species (M. triloba, M. hypoleuca, M. hosei, M. hulletti) that host plants also benefit considerably from ant-occupation. Ants do not contribute to the nutrient demands of their host plant, they do, however, protect it against herbivores and plant competition. Cleaning behaviour of the. ants results in the removal of potential herbivores already in their earliest developmental stages. Strong aggressiveness and a mass recruiting system enable the ants to defend the host plant against many herbivorous insects. This results in a significant decrease in leaf damage due to herbivores on ant-occupied compared to ant-free myrmecophytes as well as compared to non-myrmecophytic M acaranga species. Most important is the ants' defense of the host plant against plant competitors, especially vines, which are abundant in the well-lit pioneer habitats where Macaranga grows. Ants bite off any foreign plant part coming into contact with their host plant. Both ant-free myrmecophytes and non-myrmecophytic Macaranga species had a significantly higher incidence of vine growth than specimens with active ant colonies. This may be a factor of considerable importance allowing Macaranga plants to grow at sites of strongest competition.

Many plants are defended indirectly by mutualistic animals. In this field study, we investigated the efficacy of indirect antiherbivore defence in symbiotic and non-symbiotic ant-plant associations in three Macaranga species. We tested whether obligate interactions are more effective than facultative ones by comparing ant-free plants, or parts of plants, with untreated controls. All three species gained significant protection from the ants' presence. The efficacy of defence was higher in the obligate associations represented by M. triloba and M. hosei than in the facultative interaction (M. tanarius). After 40 days of ant exclusion, missing leaf area amounted to 1.7% in M. hosei (compared to 0.2% in untreated, ant-defended controls), 2.6% in M. triloba (controls 1.2%) and 4.2% in M. tanarius (controls 3.2%). In a long-term study of M. triloba and M. hosei, ant protection was orders of magnitude higher than in the short-term results. Short-term experiments obviously are unsuited to obtaining a realistic picture of the long-term efficacy of antiherbivore defence. Within 1 year, ant-free plants lost, on average, between 70% (M. hosei) and 80% (M. triloba) of their total leaf area. Both species appear to require their mutualistic ants for survival. Defence via symbiotic ants is obviously a very effective form of antiherbivore protection. Ants are highly mobile and defend preferentially young, vulnerable leaves and shoot tips, and they fulfil several functions which normally have to be provided by different chemical substances. This may be a general benefit of indirect plant defence, which makes use of "animal-specific" traits rather than intrinsic plant properties.

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