K. Eduard Linsenmair

Altitudinal distribution of leaf litter ants along a transect in primary forests on Mount Kinabalu, Sabah, Malaysia

The ant communities of the leaf litter were studied along an elevational gradient on Mount Kinabalu in primary rain forest systems ranging from dipterocarp hill forest to dwarf forest of the highest altitudes (560, 800, 1130, 1360, 1530, 1740, 1930, 2025, 2300, 2600 m a.s.l.). The litter ant fauna along the gradient included 283 species of 55 genera. The number of ant species in the leaf litter decreased exponentially without evidence of a peak in species richness at mid-elevations. This result is in contrast to many findings on altitudinal gradients in ants and other animal groups. Most ant species have a very limited altitudinal range leading to high turnover values when comparing communities of different altitudes. Of the ant species, 74% were even restricted to one site. As evident from this study, altitudinal ranges of species are very narrow. Elevational gradients are therefore extremely species-rich and might serve as a prime example of hot spots of biodiversity. This fact is of great concern when implementing conservation strategies.

Diversity of ant-plant interactions : protective efficacy in Macaranga species with different degrees of ant association

The pioneer tree Macaranga in SE Asia has developed manyfold associations with ants. The genus comprises all stages of interaction with ants, from facultative relationships to obligate myrmecophytes. Only myrmecophytic Macaranga offer nesting space for ants and are associated with a specific ant partner. The nonmyrmecophytic species are visited by a variety of different ant species which are attracted by extrafloral nectaries (EFN) and food bodies. Transitional Macaranga species like M. hosei are colonized later in their development due to their stem structure. Before the colonization by their specific Crematogaster partner the young plants are visited by different ant species attracted by EFN. These nectaries are reduced and food body production starts as soon as colonization becomes possible. We demonstrated earlier that obligate ant partners can protect their Macaranga plants against herbivore damage and vine cover. In this study we focused on nonspecific interactions and studied M. tanarius and M. hosei, representing a non-myrmecophyte and a transitional species respectively. In ant exclusion experiments both M. tanarius and M. hosei suffered significantly higher mean leaf damage than controls, 37% versus 6% in M. hosei, 16% versus 7% in M. tanarius. M. tanarius offers both EFN and food bodies so that tests for different effects of these two food rewards could be conducted. Plants with food bodies removed but with EFN remaining had the lowest mean increase of herbivore damage of all experimental groups. Main herbivores on M. hosei were mites and caterpillars. Many M. tanarius plants were infested by a shootborer. Both Macaranga species were visited by various ant species, Crematogaster spp. being the most abundant. We found no evidence for any specific relationships. The results of this study strongly support the hypothesis that non-specific, facultative associations with ants can be advantageous for Macaranga plants. Food bodies appear to have lower attractive value for opportunistic ants than EFN and may require a specific dietary adaptation. This is also indicated by the fact that food body production in the transitional M. hosei does not start before stem structure allows a colonization by the obligate Crematogaster species. M. hosei thus benefits from facultative association with a variety of ants until it produces its first domatia and can be colonized by its obligate mutualist.

Stratification of ants (Hymenoptera, Formicidae) in a primary rain forest in Sabah, Borneo

The ant fauna of a rain forest in Sabah, Malaysia was sampled by using different collecting methods in three strata. In total, 524 morphospecies of ants could be distinguished. They belong to seven subfamilies and 73 genera. So far, the ant community described is the most species rich published for a primary tropical rain forest. Regarding the stratification in the forest, the leaf litter com- munity comprised as many ant species as the lower vegetation or canopy. Further- more the litter stratum had the highest generic diversity. The stratification of ants in rain forests seems to be a very strict one with the majority of species (75%) being related to only one stratum. This is in contrast to findings on the stratifica- tion of beetles in rain forests. The stratification and a radiation of some groups into vegetation and canopy, where a broad spectrum of permanent habitats exist, is responsible for the high diversity of ants in tropical rain forests.

Size does matter - effects of tropical rainforest fragmentation on the leaf litter ant community in Sabah, Malaysia

Primary tropical lowland rainforest in Sabah, Malaysia, has been largely reduced to small to medium-sized, often isolated, forest islands surrounded by a highly altered agricultural landscape. The biodiversity patterns of leaf litter ant communities were monitored in two forest fragments of differing size as well as in a contiguous forest over the course of two years. Species number and diversity in the forest isolates was significantly lower, reaching only 47.5% of the species number collected in the contiguous forest. Species density was also lower, which had led to a thinning of the ant community in the fragments. Community composition was substantially altered in the forest remnants, and an increase of tramp species with smaller fragment size was detected. These results were unexpected and alarming, as the medium-sized forest is with its 42.9 km2 a comparatively large primary forest fragment for Sabah.

Plant-attracted ants affect arthropod community structure but not necessarily herbivory

Abstract.  1. The effectiveness of ants as plant defenders is equivocal for plants that attract ants via extrafloral nectaries (EFNs).

The importance of environmental heterogeneity for species diversity and assemblage structure in Bornean stream frogs

1. Several hypotheses have been proposed to explain the structure of multi-species assemblages. Among these, abiotic environmental factors and biotic processes are often favoured. Several recent studies examining anuran communities identified environmental factors to be only of minor importance in the composition of leaf-litter and canopy assemblages in pristine forests. Instead, spatial effects and spatially structured environments were considered more important. 2. In this study, we investigated whether these findings could also be confirmed for very heterogeneous stream habitats in the primary rainforest of the Ulu Temburong National Park, Brunei Darussalam. We thus investigated anuran assemblage compositions on 50 stream sites with regard to environmental and spatial influences. 3. Cross-product correlations indicated that both factors (spatial and environmental parameters) determined assemblage composition of anurans. Environment itself may be spatially structured, yet this interrelation did not contribute to the explainable variation of frog community compositions within the study area. 4. Detailed analyses of the environmental parameters with nonmetric multidimensional scaling revealed that community structure was mostly affected by three major environmental characters: stream turbidity, river size and the density of understorey vegetation. Based on these habitat characteristics, we assigned species to three distinct habitat guilds. 5. The results underline the importance of riparian habitat heterogeneity in pristine forests in structuring anuran assemblages. We conclude that different anuran assemblages, that is, leaf litter, canopy and stream communities, follow different assemblage rules and thus are not directly comparable.

Distribution and abundance of plants with extrafloral nectaries in the woody flora of a lowland primary forest in Malaysia

The first data on the taxonomic distribution and abundance of woody plants with extrafloral nectaries (EFN) from SE Asia are reported. The species richness and frequency of woody angiosperm plants with extrafloral nectaries was studied in the Pasoh Forest Reserve, a primary lowland forest in Peninsular Malaysia. EFN were present on 12.3% of the 741 species surveyed. 91 plant species belonging to 47 genera and 16 families were found to have EFN. Euphorbiaceae, Dipterocarpaceae, Rosaceae, Leguminosae, and Ebenaceae were the families most frequently bearing EFN whereas EFN were rare in the more primitive subclasses of the Magnoliopsida and common in the more advanced taxa (Dilleniidae, Rosidae, Asteridae). Most common were flattened glands associated with the leaf blade. A comparison with data from the Neotropics showed a great similarity in regard to the taxonomic distribution of plants with EFN. EFN-plants occupied 19.3% of the cover of the Pasoh Forest 50 ha plot.Species with EFN were under-represented among shrubs and trees of the understorey. We found an increase in the number of species with EFN as well as in their cover from the understorey to the canopy emergents. EFN were found more often among the abundant species (species with n>500/50 ha). Percentage occurrence and cover of EFN-bearing plants in the 50 ha plot of primary forest was lower than recorded for secondary habitats in tropical areas. At present, in the core zone of the Pasoh Forest Reserve which has been investigated only few species known to indicate disturbance occur. Therefore most of the recorded EFN-species cannot be regarded as secondary forest plants.The interactions between ants and EFN-bearing plants appear to be rather facultative and nonspecific. In Pasoh we found 28 ant species from seven genera visiting EFN. Most of the EFN-associated ants belonged to the subfamily Myrmicinae while Ponerines were rare, a pattern which was also reported for the Neotropics.

The Importance of Primary Tropical Rain Forest For Species Diversity: An Investigation Using Arboreal Ants as an example

The arboreal ant communities of a primary lowland rain forest and three differently disturbed forests that lie close together forming an anthropogenic disturbance gradient were collected with insecticide fogging. Combined samples from all trees (87 foggings) comprised 153,504 ants sorted to 331 morphospecies. The primary forest ant fauna was characterized by high species richness and 53 foggings were necessary to collect communities representatively. Another 63 species of ants were found in the disturbed forests indicating a large regional species pool that might exceed 420 species of arboreal ants. Anthropogenic disturbance caused a change in the taxonomic composition, diversity and structure of ant communities. Community size was a predictor of species richness in the severely disturbed forest types but not in the old secondary or primary forest. Ant abundance had declined significantly in the disturbed forests and only 10% of the primary forest’s species were collected in the most disturbed forest type. In each of the secondary forests a change in the frequency distribution of species was observed and a small number of species had gained numerical dominance. Analysis of species associations indicates that the strength of species interactions changed with the degree of forest disturbance. These changes were still clearly recognizable after 40 years of forest regeneration despite optimal conditions for colonization from the adjacent primary forest, demonstrating that the time scale needed for forest recovery after anthropogenic disturbance is very long.

Effects of Habitat Disturbance can be Subtle Yet Significant: Biodiversity of Hawkmoth-Assemblages (Lepidoptera: Sphingidae) in Southeast-Asia

Sphingid biodiversity was compared in a large number of light-trapping samples on Borneo and elsewhere in the Indo-Australian tropics, using our own quantitative light-trapping samples supplemented by other collectors’ published and unpublished data. No effects of anthropogenic habitat disturbance on the within-habitat diversity (measured as Fisher’s α) were observed, but the faunal composition of assemblages differs significantly under varying degrees of disturbance. Altitude, year of sampling and sampling regime (full night vs. part of the night) were identified as additional parameters that influence the composition of local samples. The frequency of subfamilies in samples varies under different disturbance regimes: Smerinthinae decline along a gradient from primary habitats to heavily disturbed sites, while Macroglossinae show the reverse trend. Connections between the reactions of subfamilies to disturbance and altitude and potential life-history differences between the subfamilies are discussed. Hypothetically, capital breeding Smerinthinae might be commoner and more speciose in stable primary habitats, whereas many income breeding Macroglossinae are probably adapted to thrive in ephemeral, disturbed habitats. Furthermore, we show that estimates from local samples fall short of the total known species richness of Borneo by about 10%.

The influence of anthropogenic disturbances on the structure of arboreal arthropod communities

In the framework of our research, aimed at understanding the processes structuring tropical arthropod communities, we investigated the changes occurring in tree crown communities of forests of different disturbance levels. These were a mixed dipterocarp primary lowland rain forest in Kinabalu National Park (in Sabah, Malaysia) and, some kilometres away, three forests of regeneration periods 5, 15, and 40 years following a clear-cut. These disturbed forest sites were adjacent to one another and merged into mature forest. From each forest at least ten individuals of one tree species were sampled using the fogging method. In the primary forest relative proportions of some arthropod taxa differed on the ordinal and familial level significantly within trees. The dominance of Formicidae was characteristic as was the almost complete lack of less mobile arthropods such as Lepidoptera larvae. In the five-year- old forest, differences in relative proportions among most taxa had almost disappeared. Formicidae abundances had declined drastically which coincided with an increase of Lepidoptera larvae. With progressing forest succession, arthropod communities increasingly converged on the pattern of primary forest, and total ant abundance as well as diversity increased significantly. Ant communities in the most disturbed forest were of low structural complexity, and to a large degree predictable in species arrangement, but became more and more unpredictable as the complexity of the forest increased. Several species of Coleoptera and non-formicine Hymenoptera occurred in high numbers in the youngest forest, contrasting with the mature forest where all species were typically rare. These changes may indicate a change in the structuring mechanisms from predominantly deterministic processes in disturbed forests to stochastic processes in mature forest.