Thresholds for adding degraded tropical forest to the conservation estate.

Logged and disturbed forests are often viewed as degraded and depauperate environments compared with primary forest. However, they are dynamic ecosystems1 that provide refugia for large amounts of biodiversity2,3, so we cannot afford to underestimate their conservation value4. Here we present empirically defined thresholds for categorizing the conservation value of logged forests, using one of the most comprehensive assessments of taxon responses to habitat degradation in any tropical forest environment. We analysed the impact of logging intensity on the individual occurrence patterns of 1,681 taxa belonging to 86 taxonomic orders and 126 functional groups in Sabah, Malaysia. Our results demonstrate the existence of two conservation-relevant thresholds. First, lightly logged forests (<29% biomass removal) retain high conservation value and a largely intact functional composition, and are therefore likely to recover their pre-logging values if allowed to undergo natural regeneration. Second, the most extreme impacts occur in heavily degraded forests with more than two-thirds (>68%) of their biomass removed, and these are likely to require more expensive measures to recover their biodiversity value. Overall, our data confirm that primary forests are irreplaceable5, but they also reinforce the message that logged forests retain considerable conservation value that should not be overlooked.

The first record of the parasitic myrmecophilous caterpillar Liphyrabrassolis (Lepidoptera, Lycaenidae) inside Asian weaver ant (Oecophyllasmaragdina) nests in oil palm plantations.

Asian weaver ants (Oecophyllasmaragdina) are an important biocontrol agent in agricultural habitats. We conducted surveys in oil palm plantations in Riau, Indonesia for an obligate myrmecophilous butterfly larvae, Liphyrabrassolis (Lepidoptera, Lycaenidae), that is known to consume weaver ant larvae in other habitat types. We found L.brassolis larvae in five of the twenty nests surveyed, with larval presence not being related to weaver ant nest size. We also observed L.brassolis larvae in a weaver ant mass rearing facility. This is the first report of L.brassolis from oil palm plantations and may have implications for the use of weaver ants as biological control agents.

Limited contributions of plant pathogens to density-dependent seedling mortality of mast fruiting Bornean trees.

Fungal pathogens are implicated in driving tropical plant diversity by facilitating strong, negative density-dependent mortality of conspecific seedlings (C-NDD). Assessment of the role of fungal pathogens in mediating coexistence derives from relatively few tree species and predominantly the Neotropics, limiting our understanding of their role in maintaining hyper-diversity in many tropical forests. A key question is whether fungal pathogen-mediated C-NDD seedling mortality is ubiquitous across diverse plant communities. Using a manipulative shadehouse experiment, we tested the role of fungal pathogens in mediating C-NDD seedling mortality of eight mast fruiting Bornean trees, typical of the species-rich forests of South East Asia. We demonstrate species-specific responses of seedlings to fungicide and density treatments, generating weak negative density-dependent mortality. Overall seedling mortality was low and likely insufficient to promote overall community diversity. Although conducted in the same way as previous studies, we find little evidence that fungal pathogens play a substantial role in determining patterns of seedling mortality in a SE Asian mast fruiting forest, questioning our understanding of how Janzen-Connell mechanisms structure the plant communities of this globally important forest type.

Ant mosaics in Bornean primary rain forest high canopy depend on spatial scale, time of day, and sampling method.

BACKGROUND: Competitive interactions in biological communities can be thought of as giving rise to "assembly rules" that dictate the species that are able to co-exist. Ant communities in tropical canopies often display a particular pattern, an "ant mosaic", in which competition between dominant ant species results in a patchwork of mutually exclusive territories. Although ant mosaics have been well-documented in plantation landscapes, their presence in pristine tropical forests remained contentious until recently. Here we assess presence of ant mosaics in a hitherto under-investigated forest stratum, the emergent trees of the high canopy in primary tropical rain forest, and explore how the strength of any ant mosaics is affected by spatial scale, time of day, and sampling method. METHODS: To test whether these factors might impact the detection of ant mosaics in pristine habitats, we sampled ant communities from emergent trees, which rise above the highest canopy layers in lowland dipterocarp rain forests in North Borneo (38.8-60.2 m), using both baiting and insecticide fogging. Critically, we restricted sampling to only the canopy of each focal tree. For baiting, we carried out sampling during both the day and the night. We used null models of species co-occurrence to assess patterns of segregation at within-tree and between-tree scales. RESULTS: The numerically dominant ant species on the emergent trees sampled formed a diverse community, with differences in the identity of dominant species between times of day and sampling methods. Between trees, we found patterns of ant species segregation consistent with the existence of ant mosaics using both methods. Within trees, fogged ants were segregated, while baited ants were segregated only at night. DISCUSSION: We conclude that ant mosaics are present within the emergent trees of the high canopy of tropical rain forest in Malaysian Borneo, and that sampling technique, spatial scale, and time of day interact to determine observed patterns of segregation. Restricting sampling to only emergent trees reveals segregatory patterns not observed in ground-based studies, confirming previous observations of stronger segregation with increasing height in the canopy.

Logging cuts the functional importance of invertebrates in tropical rainforest.

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests.

Experimentally testing and assessing the predictive power of species assembly rules for tropical canopy ants.

Understanding how species assemble into communities is a key goal in ecology. However, assembly rules are rarely tested experimentally, and their ability to shape real communities is poorly known. We surveyed a diverse community of epiphyte-dwelling ants and found that similar-sized species co-occurred less often than expected. Laboratory experiments demonstrated that invasion was discouraged by the presence of similarly sized resident species. The size difference for which invasion was less likely was the same as that for which wild species exhibited reduced co-occurrence. Finally we explored whether our experimentally derived assembly rules could simulate realistic communities. Communities simulated using size-based species assembly exhibited diversities closer to wild communities than those simulated using size-independent assembly, with results being sensitive to the combination of rules employed. Hence, species segregation in the wild can be driven by competitive species assembly, and this process is sufficient to generate observed species abundance distributions for tropical epiphyte-dwelling ants.

An ant-plant by-product mutualism is robust to selective logging of rain forest and conversion to oil palm plantation.

Anthropogenic disturbance and the spread of non-native species disrupt natural communities, but also create novel interactions between species. By-product mutualisms, in which benefits accrue as side effects of partner behaviour or morphology, are often non-specific and hence may persist in novel ecosystems. We tested this hypothesis for a two-way by-product mutualism between epiphytic ferns and their ant inhabitants in the Bornean rain forest, in which ants gain housing in root-masses while ferns gain protection from herbivores. Specifically, we assessed how the specificity (overlap between fern and ground-dwelling ants) and the benefits of this interaction are altered by selective logging and conversion to an oil palm plantation habitat. We found that despite the high turnover of ant species, ant protection against herbivores persisted in modified habitats. However, in ferns growing in the oil palm plantation, ant occupancy, abundance and species richness declined, potentially due to the harsher microclimate. The specificity of the fern-ant interactions was also lower in the oil palm plantation habitat than in the forest habitats. We found no correlations between colony size and fern size in modified habitats, and hence no evidence for partner fidelity feedbacks, in which ants are incentivised to protect fern hosts. Per species, non-native ant species in the oil palm plantation habitat (18 % of occurrences) were as important as native ones in terms of fern protection and contributed to an increase in ant abundance and species richness with fern size. We conclude that this by-product mutualism persists in logged forest and oil palm plantation habitats, with no detectable shift in partner benefits. Such persistence of generalist interactions in novel ecosystems may be important for driving ecosystem functioning.

The first record of a fly of the family Milichiidae (Diptera) interacting with an ant of the genus Polyrhachis Smith, 1857 (Hymenoptera: Formicidae).

Flies in the family Milichiidae are often myrmecophilic. We document the first record of a fly from this family interacting with an ant of the genus Polyrhachis. In lowland riparian rainforest in Sabah, Malaysia, we observed a female of the genus Milichia following an ant of the species of P.illaudata, and repeatedly attempting to make close contact. Our observation suggests that the dipteran may have been attempting to feed kleptoparasitically from the Polyrhachis worker, since members of this ant genus often feed on liquid carbohydrate-rich food resources. This is the first time an interaction has been observed between a fly of this family and an ant of this widespread old world tropical genus.

Establishing the evidence base for maintaining biodiversity and ecosystem function in the oil palm landscapes of South East Asia.

The conversion of natural forest to oil palm plantation is a major current threat to the conservation of biodiversity in South East Asia. Most animal taxa decrease in both species richness and abundance on conversion of forest to oil palm, and there is usually a severe loss of forest species. The extent of loss varies significantly across both different taxa and different microhabitats within the oil palm habitat. The principal driver of this loss in diversity is probably the biological and physical simplification of the habitat, but there is little direct evidence for this. The conservation of forest species requires the preservation of large reserves of intact forest, but we must not lose sight of the importance of conserving biodiversity and ecosystem processes within the oil palm habitat itself. We urgently need to carry out research that will establish whether maintaining diversity supports economically and ecologically important processes. There is some evidence that both landscape and local complexity can have positive impacts on biodiversity in the oil palm habitat. By intelligent manipulation of habitat complexity, it could be possible to enhance not only the number of species that can live in oil palm plantations but also their contribution to the healthy functioning of this exceptionally important and widespread landscape.

The life of a dead ant: the expression of an adaptive extended phenotype.

Specialized parasites are expected to express complex adaptations to their hosts. Manipulation of host behavior is such an adaptation. We studied the fungus Ophiocordyceps unilateralis, a locally specialized parasite of arboreal Camponotus leonardi ants. Ant-infecting Ophiocordyceps are known to make hosts bite onto vegetation before killing them. We show that this represents a fine-tuned fungal adaptation: an extended phenotype. Dead ants were found under leaves, attached by their mandibles, on the northern side of saplings approximately 25 cm above the soil, where temperature and humidity conditions were optimal for fungal growth. Experimental relocation confirmed that parasite fitness was lower outside this manipulative zone. Host resources were rapidly colonized and further secured by extensive internal structuring. Nutritional composition analysis indicated that such structuring allows the parasite to produce a large fruiting body for spore production. Our findings suggest that the osmotrophic lifestyle of fungi may have facilitated novel exploitation strategies.