Temporal and Spatial Diversity and Distribution of Arboreal Carabidae (Coleoptera) in a Western Amazonian Rain Forest1

Diversity of arboreal carabid beetles was sampled by fumigation in 100 3 × 3 m stations within a 100 × 1000 m terra firme forest plot in Ecuadorian Amazonia. Nine sampling dates from January 1994 to October 1996 yielded 2329 individuals belonging to 318 species of which more than 50 percent were undescribed species. A high percentage of the species sampled were rare; the proportion that occurred once per sampling date (singletons) ranged from 50.0 to 62.5 percent. Estimates of species richness were from 82 to 282 species of arboreal carabids in the study plot on a given sampling date. Most richness values were greater than 173 species. Species accumulation curves attained asymptotes for all but one sampling date, indicating that an adequate level of sampling effort was used to characterize the diversity of carabid fauna. Total accumulation curves based on pooled data failed to reach asymptotes. There was a high turnover in species composition between sampling dates; less than 50 percent of the species between the majority of sampling dates were shared, suggesting that the total species pool may be extremely large. Although species composition changed seasonally, species richness varied little. Spatial autocorrelation analysis revealed that the structure of this species assemblage was significantly patterned at distances below 280 m. Taken together, the large percentage of undescribed species, die failure of the overall species accumulation curves to level off, and the high turnover in species composition indicate that the species diversity of carabid beetles is far higher than previously thought.     

Tree Diversity,Forest Structure and Productivity along Altitudinal and Topographical Gradients in a Species-Rich Ecuadorian Montane Rain Forest

We studied the spatial heterogeneity of tree diversity, and of forest structure and productivity in a highly diverse tropical mountain area in southern Ecuador with the aim of understanding the causes of the large variation in these parameters. Two major environmental gradients, elevation and topography, representing a broad range of climatic and edaphic site conditions, were analyzed. We found the highest species richness of trees in valleys <2100 m. Valleys showed highest values of basal area, leaf area index and tree basal area increment as well. Tree diversity also increased from ridges to valleys, while canopy openness decreased. Significant relationships existed between tree diversity and soil parameters (pH, total contents of Mg, K, Ca, N and P), and between diversity and the spatial variability of pH and Ca and Mg contents suggesting a dependence of tree diversity on both absolute levels and on the small-scale heterogeneity of soil nutrient availability. Tree diversity and basal area increment were positively correlated, partly because both are similarly affected by soil conditions. We conclude that the extraordinarily high tree species richness in the area is primarily caused by three factors: (1) the existence of steep altitudinal and topographic gradients in a rather limited area creating a small-scale mosaic of edaphically different habitats; (2) the intermingling of Amazonian lowland plant species, that reach their upper distribution limits, and of montane forest species; and (3) the geographical position of the study area between the humid eastern Andean slope and the dry interandean forests of South Ecuador.     

Diversity and composition of Arctiidae moth ensembles along a successional gradient in the Ecuadorian Andes

Andean montane rain forests are among the most species‐rich terrestrial habitats. Little is known about their insect communities and how these respond to anthropogenic habitat alteration. We investigated exceptionally speciose ensembles of nocturnal tiger moths (Arctiidae) at 15 anthropogenically disturbed sites, which together depict a gradient of forest recovery and six closed‐forest understorey sites in southern Ecuador. At weak light traps we sampled 9211 arctiids, representing 287 species. Arctiid abundance and diversity were highest at advanced succession sites, where secondary scrub or young forest had re‐established, followed by early succession sites, and were lowest, but still high, in mature forest understorey. The proportion of rare species showed the reverse pattern. We ordinated moth samples by non‐metric multidimensional scaling using the chord‐normalized expected species shared index (CNESS) index at various levels of the sample size parameter m. A distinct segregation of arctiid ensembles at succession sites from those in mature forest consistently emerged only at high m‐values. Segregation between ensembles of early vs. late succession stages was also clear at high m values only, and was rather weak. Rare species were responsible for much of the faunal difference along the succession gradient, whereas many common arctiid species occurred in all sites. Matrix correlation tests as well as exploration of relationships between ordination axes and environmental variables revealed the degree of habitat openness, and to a lesser extent, elevation, as best predictors of faunal dissimilarity. Faunal differences were not related to geographical distances between sampling sites. Our results suggest that many of the more common tiger moths of Neotropical montane forests have a substantial recolonization potential at the small spatial scale of our study and accordingly occur also in landscape mosaics surrounding nature reserves. These species contribute to the unexpectedly high diversity of arctiid ensembles at disturbed sites, whereas the proportion of rare species declines outside mature forest.     

Dung Beetle Communities and Seed Dispersal in Primary Forest and Disturbed Land in Amazonia1

Seeds from tropical fruiting trees ingested and defecated on the soil surface by primary dispersers (such as primates) are vulnerable to destruction from rodents, insects, and fungi. Burial by dung beedes as an incidental result of their feeding and nesting activities often provides these seeds with refugia from attack. To examine the effect of habitat disturbance on the dung beetle communities involved in this process, I surveyed dung beedes at three sites in the Amazon basin, in die states of Parí, Amazonas, and Rondónia, Brazil. Through principal component analysis on measurements of size and behavioral characters of beedes, I determined the relative quality of beetle species as seed dispersers (dispersal defined as horizontal or vertical movement of seeds) and ranked them into seed dispersal guilds. I used correspondence analysis to examine in what habitats (primary forest or varying degrees of disturbed habitat) these guilds were abundant. Most guilds decreased with increasing habitat disturbance, but one guild made up of large nocturnal burrowers (primarily Dichotomius) became more abundant with increasing disturbance (up to the level of highest disturbance surveyed), at which point all dung beede species became scarce. Clear–cuts had lower species richness, lower abundance, and lower biomass dian forested areas. These results imply that clear–cutting is detrimental to all seed dispersal dynamics in tropical rain forests, but that some levels of disturbance allow enough disperser activity to preserve this ecosystem function.     

The Structure of Ground Beetle Assemblages (Coleoptera: Carabidae) at Fruit Falls of Melastomataceae Trees in a Brazilian Terra Firme Rain Forest1

Although most Carabidae are recognized as predacious, recent studies have shown that some tropical rain forest Carabidae use the small seeds of fallen fruit from some tree species as a key food resource (particularly figs [Moraceae] but also those from other families). Assemblages of carabid beetles (Coleoptera: Carabidae) at fruit falls of the Melastomataceae tree genera, Bellucia, Loreya, and Miconia, were studied in a terra firme rain forest near Manaus (Amazonia, Brazil) from February 1992 to August 1996. Of 5483 specimens of 14 carabid species collected at 53 fruit falls, 98.2 percent of the collected beetles represented the seed-feeding genus Notiobia. Four of the 7 Notiobia species were found to reproduce at Melastomataceae fruit falls. The most abundant species, N. glabrata, was found to be a Melastomataceae seed generalise while N. maxima seemed to prefer the seeds of Bellucia, and N. aulica and Af. umbrifera the seeds of Miconia. The presence of larvae indicated reproduction of N. glabrata at fruit falls of all three tree genera and reproduction of N. maxima at those of Bellucia. Fruit falls of different species and genera of Melastomataceae appeared to occur at different times of die year and not continuously throughout the year. The seed-feeding Notiobia species, which specialize on Melastomataceae, used fruit falls of figs as migratory “stepping stones” during periods when there were no Melastomataceae fruit falls. The comparable suite of fig seed-feeding specialist species of Notiobia did not appear to do the reverse and seemed to avoid Melastomataceae fruit falls. The carabid assemblage at fig fruit falls was more diverse than the assemblage at those of Melastomataceae. Among the Melastomataceae, carabid species richness was highest at Miconia fruit falls and lowest at Loreya fruit falls. The three most common predatory species caught at Ficus fruit falls were also caught at those of Melastomataceae and appeared to reproduce at both fruit fall types. The proportion of the carabid assemblage accounted by predatory species was much lower at Melastomataceae (1.8%) than at fig fruit falls (8%).     

Edaphic and Floristic Variation within a 1‐ha Plot of Lowland Amazonian Rain Forest1

Several studies in lowland tropical rain forests have documented effects of local‐scale topographic variation on plant species distribution and abundance patterns. Few studies have compared the distribution patterns of more than one plant group, however, and even fewer have related these to measured physical and chemical soil characteristics. Here, we document such soil characteristics within a square 1‐ha plot in Amazonian Ecuador, and compare them to the distribution patterns of terrestrial pteridophytes, angiosperm ground herbs, and palms. Substantial variation in soil properties was found within the plot. The three plant groups showed highly correlated floristic patterns within the 1‐ha plot even after the effect of geographical distances had been taken into account. Mantel tests yielded significant correlations between edaphic patterns, as measured by distances in various soil and topographic characteristics, and floristic patterns. For all three plant groups, differences in elevation within the plot were highly correlated with floristic distances, and for terrestrial pteridophytes and palms, distances in soil calcium content and sand content were also important. Our results resembled those obtained at wider spatial scales with the same plant groups, which indicates that soil factors may play an important role for distribution and beta diversity of plants, even at the local scale.     

Factors Affecting the Small Mammal Community Inside and Outside Katavi National Park, Tanzania1

Small mammal trapping showed that a minimum of 12 species live in and around Katavi National Park in western Tanzania. Species richness and abundance were greater outside the park than inside, extending results of an earlier study. Species richness and abundance were not significantly associated with vegetation cover or plant biomass but were negatively correlated with seed diversity, seed biomass, and possibly small carnivore abundance. Unfortunately, small mammal diets are unknown in this ecosystem and it can be argued that carnivores are unlikely to reduce species diversity, casting doubt on the importance of these variables. While the factors causing small mammals to fare poorly in this protected area are not yet understood, reduced food availability remains the best candidate.     

Association of Ant Predators and Edaphic Conditions with Termite Diversity in an Amazonian Rain Forest

Predation is a key determinant of prey community structure, but few studies have measured the effect of multiple predators on a highly diverse prey community. In this study, we asked whether the abundance, species richness, and species composition of a species‐rich assemblage of termites in an Amazonian rain forest is more strongly associated with the density of predatory ants or with measures of vegetation, and soil texture and chemistry. We sampled termite assemblages with standardized hand‐collecting in 30 transects arranged in a 5 km × 6 km grid in a terra firme Amazonian rain forest. For each transect, we also measured vegetation structure, soil texture, and soil phosphorus, and estimated the density of predatory ants from baits, pitfall traps, and Winkler samples. Seventy‐nine termite species were recorded, and the total density of predatory ants was the strongest single predictor of local termite abundance (r = ?0.66) and termite species richness (r = ?0.44). In contrast, termite abundance and species richness were not strongly correlated with edaphic conditions (¦r¦ < 0.01), or with the density of non‐predatory ants (r_abund = ?0.27; r_s = ?0.06). Termite species composition was correlated with soil phosphorus content (r = 0.79), clay content (r = ?0.75), and tree density (r = ?0.42). Assemblage patterns were consistent with the hypothesis that ants collectively behaved as generalist predators, reducing total termite abundance, and species richness. There was no evidence that ants behaved as keystone predators, or that any single termite species benefited from the reduction in the abundance of potential competitors.     

Broad Scale Distribution of Ferns and Lycophytes along Environmental Gradients in Central and Northern Amazonia,Brazil

Establishing which factors determine species distributions is of major relevance for practical applications such as conservation planning. The Amazonian lowlands exhibit considerable internal heterogeneity that is not apparent in existing vegetation maps. We used ferns as a model group to study patterns in plant species distributions and community composition at regional and landscape scales. Fern species composition and environmental data were collected in 109 plots of 250 × 2 m distributed among four sites in Brazilian Amazonia. Interplot distances varied from 1 to ca 670 km. When floristically heterogeneous datasets were analyzed, the use of an extended Sørensen dissimilarity index rather than the traditional Sørensen index improved model fit and made interpretation of the results easier. Major factors associated with species composition varied among sites, difference in cation concentration was a strong predictor of floristic dissimilarity in those sites with pronounced heterogeneity in cation concentration. Difference in clay content was the most relevant variable in sites with uniform cation concentrations. In every case, environmental differences were invariably better than geographic distances in predicting species compositional differences. Our results are consistent with the ideas that: (1) the relative predictive capacity of the explanatory variables depend on the relative lengths of the observed gradients; and (2) environmental gradients can be hierarchically structured such that gradients occur inside gradients. Therefore, site‐specific relationships among variables can mask the bigger picture and make it more difficult to unravel the factors structuring plant communities in Amazonia.     

Ecology and Distribution of Melastomataceae in African Rain Forest Inselbergs1

The ecology and distribution of thirty‐seven Melastomataceae species found on 29 inselbergs in the rain forest of Equatorial Guinea, Gabon, and Cameroon are described. The relative contributions of climatic determinism, and stochastic and historical processes to the present distribution of the species amongst the study sites are investigated. Inselbergs are rock outcrops enduring very harsh edaphic and microclimatic conditions, forming “xeric islands” in the rain forest, where natural grasslands develop. For each Melastomataceae species, the habitats where they were collected on the inselbergs and in other biotopes were listed, and their phytogeographical distribution was established. Three species are restricted to rock outcrops. Most species are distributed inside the Guineo‐Congolian Region. Three species have disjunct distributions on rain forest inselbergs and have their main distribution area outside the forested zone. The distribution of the species amongst the inselbergs is correlated to the altitude and the annual rainfall of the sites, and to inselberg geographical location. This correlation of the species distribution with the geographical location of the inselberg is attributed to the spatial autocorrelation of environmental variables and to the combined effects of the history of past vegetation changes due to climatic fluctuations and dispersal limitation.     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

Effects of Selective Logging on Populations of Two Tropical Understory Herbs in an Amazonian Forest1

This study examined the effects of (1) rime elapsed after logging and (2) logging intensity on the total, juvenile, and adult densities, and on reproduction of Calathea altissima and Ischnosiphon amuma (Matantaceae), two monocot herb species in the understory of Central Amazonian forests. Logging was carried out experimentally at various intensities in eight plots (4 ha each) during 1987 and three plots in 1993. Five plots were left as experimental controls. In 1998, total and adult densities of the two species were greater in the plots logged 11 years before than in controls. For I. arouma, total and adult densities were also higher in the plots logged 5 years before; however, juvenile densities were less affected. The intensity of logging influenced adult density of both species but not total or juvenile densities. Densities of reproductive individuals of both species were higher in the logged areas and increased with logging intensity. The effects of time after logging and logging intensity on reproduction were indirect due to the greater number of adult plants in those areas. The observed effects were probably mediated by changes in canopy cover in logged areas. Greater light intensities in logging gaps, roads, and their margins may lead to increased reproduction, recruitment, growth, and survival, which in turn can lead to increased plant density. These results indicate that logging has long–term effects on understory plant populations.     

Seasonal Change of Species Diversity Patterns of Non‐volant Small Mammals along Three Subtropical Elevational Gradients

Our understanding of geographic patterns of species diversity and the underlying mechanisms is increasing rapidly, whereas the temporal variation in these patterns remains poorly understood. We examined the seasonal species richness and species turnover patterns of non‐volant small mammals along three subtropical elevational gradients in southwest China. Small mammal diversity was surveyed in two seasons (early wet season and late wet season) using a standardized sampling protocol. The comparison of species richness patterns between two seasons indicated a temporal component in magnitude and shape, with species richness at high elevations clearly increased during the late wet season. Species richness demonstrated weak correlations with modelled temperature and precipitation. The elevational pattern of species turnover measured by Chao‐Sørenson similarity index also changed seasonally, even though the temporal pattern varied with scale. Species turnover between neighboring elevations at high elevations was slower in the late wet season. Meanwhile, there was an acceleration of species turnover along the whole range of the gradient. The seasonal change in species diversity patterns may be due to population‐level increases in abundance and elevational migration, whereas seasonal variation in factors other than temperature and precipitation may play a greater role in driving seasonal diversity patterns. Our study strongly supports the seasonality in elevational patterns of small mammal diversity in subtropical montane forests. Thus it is recommended that subsequent field surveys consider temporal sampling replicate for elevational diversity studies.