An analysis was conducted of floristic patterns contained in 48 1-ha tree plots distributed at 29 sites in seven neotropical countries, with a primary emphasis on the Amazonian region. Analyses were made with family level data, using detrended correspondence analysis and multidimensional scaling to generate two-dimensional ordinations. Dissimilarity values for all pairs of plots were then used to compare forest composition at both local (flooded vs unflooded forests) and regional scales (e.g., western vs central vs eastern Amazonia). The predominate family of trees in a large majority of Amazonian and Guianan forests (by number of stems) is either Palmae or Leguminosae (sensu latu), followed by Moraceae and Euphorbiaceae. The forests of western Amazonia are particularly rich in palms, Moraceae, and Myristicaceae, whereas those of eastern Amazonia and the Guianas are rich in Lecythidaceae and Chrysobalanaceae. Dissimilarity between sites increases with distance for both flooded and unflooded forests. The tree communities of flooded and unflooded forests within a region tended to resemble one another more closely than forests of either type resembled the homologous forests of the adjoining regions. Within Amazonia the edaphic properties of each region and its geological history are tightly interrelated. It is therefore difficult to distinguish between evolutionary and ecological interpretations of the results.

Although there is increasing interest in the effects of habitat disturbance on community attributes and the potential consequences for ecosystem functioning, objective approaches linking biodiversity loss to functional loss are uncommon. The objectives of this study were to implement simultaneous assessment of community attributes (richness, abundance and biomass, each calculated for total-beetle assemblages as well as small- and large-beetle assemblages) and three ecological functions of dung beetles (dung removal, soil perturbation and secondary seed dispersal), to compare the effects of habitat disturbance on both sets of response variables, and their relations. We studied dung beetle community attributes and functions in five land-use systems representing a disturbance gradient in the Brazilian Amazon: primary forest, secondary forest, agroforestry, agriculture and pasture. All response variables were affected negatively by the intensification of habitat disturbance regimes, but community attributes and ecological functions did not follow the same pattern of decline. A hierarchical partitioning analysis showed that, although all community attributes had a significant effect on the three ecological functions (except the abundance of small beetles on all three ecological functions and the biomass of small beetles on secondary dispersal of large seed mimics), species richness and abundance of large beetles were the community attributes with the highest explanatory value. Our results show the importance of measuring ecological function empirically instead of deducing it from community metrics.

Abstract. 1. The role of several factors that affect the composition of the dung beetle assemblages in an Amazonian rainforest was quantified, together with the effect of these factors on the role that dung beetles play as secondary seed dispersers.2. A total of 61 dung beetle species was captured during 3360 h of trapping. During nocturnal trapping periods, more dung beetles, of larger mean size, and more species were captured per trap than during diurnal trapping periods.3. During the rainy season, more dung beetle species were captured per trap than during the dry season, but the number of individuals and their mean size did not vary between seasons.4. Bait size had a significant effect on the mean number of beetles and mean number of species but not on mean beetle size. As bait size increased from 5, 10, 25, to 50 g, more beetles and more species were captured per trap.5. Between 6 and 73% of plastic beads, used as seed mimics, were buried by dung beetles at depths that ranged from 0.5 to 7 cm. Both the proportion of beads buried and burial depth decreased with increasing bead size, and increased with increasing amounts of dung surrounding each bead (5, 10, and 25 g).6. The proportion of buried seeds for three species varying in size between 5 and 27 mm, increased with increasing dung beetle size, using beetles of seven sizes, varying between 10 and 25 mm.7. Seeds surrounded by dung were buried more often and more deeply when placed on the forest floor during the late afternoon than when placed during the early morning. Seeds were also buried more often when placed on the forest floor during the rainy season than when placed during the dry season, but season had no effect on burial depth.8. Forests in Central Amazonia hold a rich dung beetle community that plays an active role in secondary seed dispersal, and consequently in plant regeneration. The interaction between seeds and beetles is complex because it is affected by many factors.

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung ( n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5-20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.

Andresen, E. 2003. Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration. -Ecography 26: 87 -97.I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.E. Andresen (andresen@oikos.unam.mx),

This paper reviews the current knowledge on secondary seed dispersal by dung beetles and its effect on plant regeneration in tropical rain forests. In the first section, the intrinsic factors affecting the outcome of the seed-beetle interaction, emphasizing both the beetle's and the plant's perspective are presented. In the second section, some of the extrinsic factors that indirectly affect the outcome of the interaction are highlighted. Both sections emphasize little-known aspects of the effects of dung beetles on plant regeneration, pointing out new avenues for future research.

ABSTRACT. The study of seed fate is crucial for understanding fruit-frugivore interactions. One factor that can greatly influence the fate of seeds dispersed through mammal defecation, is the dung that accompanies the seeds. Dung attracts dung beetles and rodents; the former eat and bury dung, the latter feed on seeds. In this study the fate of Micropholis guyanensis subsp. guyanensis seeds surrounded by no dung and by 5, 10 and 25 g of howler monkey dung was followed until seedling establishment. The depths at which dung beetles bury the seeds were measured, and the effect of burial depth on seedling emergence was determined. Although initial seed fates differed among treatments, the same percentage of seedlings established from seeds without faecal material, and from seeds with 5, 10 and 25 g of dung. However, a significantly higher proportion of seedlings established from buried seeds when compared to seeds that remained on the surface. The percentage of seedlings establishing in a controlled germination experiment decreased significantly with increasing burial depth. The effect of dung beetle activity should be taken into consideration when assessing the role that mammal endozoochory plays on seed dispersal ecology of tropical plants. These results provide further evidence for the emerging realization that the study of secondary seed dispersal and post-dispersal events is crucial for a more complete understanding of plant regeneration.

Dung beetles are important components of most terrestrial ecosystems. In tropical rain forests, dung beetle communities can be very rich in number of species and individuals, and they are known to be useful bioindicators of habitat disturbance. In contrast, very little is known about the organization of dung beetle communities in tropical dry forests. The aim of this study was to describe in detail the dung beetle community of a Mexican tropical dry forest and to assess the relative importance of rainfall seasonality and forest structure in affecting the temporal and spatial dynamics of this community. Dung beetles were captured with pitfall traps at the beginning of the rainy season, the middle of the rainy season, and the middle of the dry season, in two distinct forest types: deciduous forest (DF) and semideciduous forest (SDF) at the Estación de Biología Chamela. Both rainfall seasonality and forest structure affected the community organization of dung beetles. During both rainy periods, 14 species were captured, but only three during the dry season. Dung beetles captured during the dry season were only found in the SDF. When comparing the beginning and the middle of the rainy season, differences in abundance and guild structure were also observed between both periods and between forest types, but these differences were much less pronounced.