Seedling assemblages and the alternative successional pathways experienced by Atlantic forest fragments

Background: Habitat loss and fragmentation have been argued to drastically alter the composition of tree assemblages inhabiting small forest fragments but the successional trajectory experienced by such edge-affected habitats remains controversial.

Aims: Here we examine whether small fragments (3.4–91.2 ha) support seedling assemblages more similar to those in 10–70-year-old secondary forests than to those in mature forests, in order to infer to what extent fragments move toward early successional systems.

Methods: Using 59 0.1-ha plots distributed in a fragmented landscape of Brazilian Atlantic forest, we evaluated species richness and functional and taxonomic composition of seedling assemblages in 20 small forest fragments, 19 stands of secondary forest and 20 stands of mature forests in the interior of an exceptionally large fragment (ca. 3500 ha).

Results: Small fragments presented the least species-rich seedling assemblages (17.2 ± 5.7 species), followed by secondary (22.5 ± 5.3), and mature forest (28.4 ± 5.3). Small fragments had seedling assemblages with functional and taxonomic composition more similar to those in secondary than in mature forest. Small fragments had a greater relative richness and abundance of pioneer trees (ca. 40% more), vertebrate-dispersed (6–25%), and those bearing medium-sized seeds (30–70%), while large-seeded species and individuals were reduced (>50% decrement) in comparison to seedling assemblages in mature forest.

Conclusions: By comparing seedlings across a wide range of successional habitats we offer evidence that small forest fragments are experiencing an alternative successional pathway towards an early-successional system with reduced plant diversity.     

Relationships between land snail assemblage patterns and soil properties in temperate-humid forest ecosystems

Aim The objectives were to (1) analyse the combined effects of soil pH, Ca content and soil moisture on total density and species richness of land snails in forest ecosystems, (2) explore relationships between the quantitative composition of snail assemblages and habitat characteristics, (3) investigate the relationships between soil pH and density of some of the most frequent species, and (4) compare the data with those from studies conducted in other temperate‐humid regions of Europe. Location Study sites were selected from 15 landscape types including different lithologies within the area of Baden‐Württemberg (35,000 km²), SW Germany. Methods Snails were recorded quantitatively from 83 study sites, with four plots representing a total of 0.25 m² per site. Topsoil samples from each site were analysed for pH, exchangeable Ca, and Ca content of carbonates. Three categories of soil moisture (dry, intermediate and wet) were established and defined according to the (climatic) water balance. Numbers of individuals and species were brought in relation to soil moisture and soil pH. Cluster analyses were conducted to identify groups of quantitatively similar snail species assemblages. Results Topsoil pH (2.7–7.5) and soil Ca contents were closely correlated. On dry soils, total snail density and species richness are generally low and do not change with pH, but clearly increase with increasing pH on intermediate moisture soils and on wet soils. On the latter, numbers of individuals and species are generally much higher compared with intermediate moisture sites at the same value of soil pH. Changes of density in relation to soil pH vary between species. Depending on the species, density increases only in the lower or only in the higher range of pH, is not related to pH, or decreases with increasing pH. Furthermore, these patterns vary within the same species depending on the region. This became evident from comparisons with other studies, particularly between sites in SW Germany and southern Scandinavia. From cluster analyses, subgroups of snail assemblages of high quantitative similarity were identified. Group formation is explained by soil pH to some extent, and one subgroup showed a connection with coniferous woodland sites on acidic soils. No further environmental factors available from our data could explain the clustering of snail assemblages more detailed. Main conclusions Soil moisture is the strongest determinant of snail density and species richness at undisturbed woodland sites, but effects of soil moisture and soil pH on these patterns are closely interrelated on intermediate moisture soils and wet soils. However, the quantitative species composition of the land snail assemblages is related to soil properties to a lower degree than snail density and species richness, and other habitat characteristics such as vegetation or litter quality, can be important for species dominance in addition.     

Abundance and cover of ground herbs in an Amazonian rain forest

In a 1-ha plot in an unflooded moist tropical forest in Reserva de Producción Faunística Cuy abeno in Amazonian Ecuador, 96 species of vascular herbaceous plants were found rooted in the ground; they were all perennials; 25 species were pteridophytes, representing 11 families, 71 species (14 families) were angiosperms. Araceae, Marantaceae, and Poaceae were the most important angiosperm families. The total abundance of the ground herbs was 10 960 individuals, the total cover was 250 m², or 2.5% of the 1-ha plot. Species of ground herbs exhibit two major life-form strategies: the obligate terrestrial species (59%) are restricted to the ground; the facultative terrestrial species (41 %) have climbing and epiphytic individuals as well. Difference in life-form strategy as well as difference in edaphic specialization along a topographic gradient are two factors that may enhance the number of coexisting species within the 1-ha sample plot.     

Tree species richness in primary and old secondary tropical forest in Singapore

If secondary succession can accumulate species rapidly, then tropical secondary forests may have an important role to play in the conservation of biodiversity. Data on the floristic composition of forest stands in the Central Catchment Nature Reserve, Singapore, have been analysed to investigate the diversity of approximately 100-year-old tropical secondary forest. Classification using TWINSPAN indicated that three floristic communities could be recognized from 59 0.2 ha plots enumerated for trees >30 cm gbh. These were two types of secondary forest, both dominated by Rhodamnia cinerea (Myrtaceae), and dryland primary forest. The secondary forest was developed on land abandoned after use for agriculture at the end of the 19th century. The 16 primary forest plots contained a total of 340 species, more than the 281 recorded from the 43 plots of the two secondary forest types combined. The mean species number per plot in the more diverse of the two secondary forests was only about 60% of the primary forest. Thus the secondary forest, despite a century or so for colonization by species and the presence of contiguous primary forest, was still significantly less diverse than primary forest areas. It is concluded that secondary forest cannot be assumed to accrete biodiversity rapidly in the tropics, and may not be of direct value in conservation. However, other indirect roles, such as providing resources for native animals, and buffering and protecting primary forest fragments may make the protection of secondary forest worthwhile.     

Effects of flooding duration on species richness, floristic composition and forest structure in river margin habitat in Amazonian blackwater floodplain forests: implications for future design of protected areas

Rivers in central Amazonia experience annual water-level fluctuations of up to 14m, flooding vast areas of adjacent forest for periods ranging from a few to 270 days per year. At different sites, variation in the duration and type of flooding results in a mosaic of habitats that includes lakes, grasslands, forests, and streams. To study the effects of flood duration on plant species richness and floristic composition, two river margin sites were surveyed on the rivers Jaú and Tarumã-Mirim. Both areas are seasonally flooded by blackwaters, and plots were made at different topographic levels (lower, middle and upper slopes). All woody plants with DBH>5cm were inventoried in five 10 × 40m plots in each of the three topographic levels, which varied in length of flood duration and mean water level. Plant species richness did not vary significantly between topographic levels, but species composition varied substantially. At both study sites, the species composition exhibited distinctive distribution patterns with respect to the three topographic levels and river site. Differences in the distribution of dominant species in both sites probably relate to the ability of species to withstand seasonal flooding, although other edaphic factors associated with the topographic levels may also be important, especially for less-dominant, locally rare, and habitat generalist species. Species composition overlap among topographic levels at the two sites was highly variable ranging from 15% to 43%. Knowledge about the complex pattern of species composition and distributions between and among topographic levels and river sites is important for the preservation of the diverse flora of the blackwater forests and for the creation of future conservation management plans and design of protected areas in this ecosystem that will maintain the biodiversity.     

Fire-mediated dieback and compositional cascade in an Amazonian forest

The only fully coupled land-atmosphere global climate model predicts a widespread dieback of Amazonian forest cover through reduced precipitation. Although these predictions are controversial, the structural and compositional resilience of Amazonian forests may also have been overestimated, as current vegetation models fail to consider the potential role of fire in the degradation of forest ecosystems. We examine forest structure and composition in the Arapiuns River basin in the central Brazilian Amazon, evaluating post-fire forest recovery and the consequences of recurrent fires for the patterns of dominance of tree species. We surveyed tree plots in unburned and once-burned forests examined 1, 3 and 9 years after an unprecedented fire event, in twice-burned forests examined 3 and 9 years after fire and in thrice-burned forests examined 5 years after the most recent fire event. The number of trees recorded in unburned primary forest control plots was stable over time. However, in both once- and twice-burned forest plots, there was a marked recruitment into the 10-20cm diameter at breast height tree size classes between 3 and 9 years post-fire. Considering tree assemblage composition 9 years after the first fire contact, we observed (i) a clear pattern of community turnover among small trees and the most abundant shrubs and saplings, and (ii) that species that were common in any of the four burn treatments (unburned, once-, twice- and thrice-burned) were often rare or entirely absent in other burn treatments. We conclude that episodic wildfires can lead to drastic changes in forest structure and composition, with cascading shifts in forest composition following each additional fire event. Finally, we use these results to evaluate the validity of the savannization paradigm.     

Separating the effects of number of individuals sampled and competition on species diversity: an experimental and analytic approach

1 Species richness typically increases with the number of individuals sampled, although many ecological processes that influence species richness are also well known to depend on density of individuals. We separated the effects of density on species richness that are due to sampling, from those due to density-dependent ecological processes such as competition or predation, by manipulating the density of an entire community.
2 A seed bank from a community of desert annual plants that occur on semi-stabilized sand dunes in Israel was collected from the field and sown in an experimental garden at a range of densities from 1/16 to eight times the natural density. The species pool observed in the lowest density plots was used as the null community, which was repeatedly sampled to calculate the species richness (and other diversity indices) in higher density plots that would be expected from sampling considerations alone. The significance of deviations of observed diversity from this expected diversity was then evaluated.
3 Both observed and expected number of species increased substantially with the experimental increase in density. However, observed species richness, the Shannon–Wiener diversity index and Simpson's diversity index were often significantly lower than that expected based on sampling considerations. The magnitude of the deviation from expected increased significantly with increasing density for richness and the Shannon–Wiener index. This provides some of the first direct experimental evidence from diverse natural assemblages that increasing competition among all the individuals in a community can lead to competitive exclusion.     

热带森林树种多度和丰富度空间分布特征研究

用巴拿马50 hm2森林动态监测样地内直径≥1 cm的树种资料,分析了该样地树种多度(个体数)和丰富度(物种数)及其方差和变异系数在6个取样尺度(5 m×5 m,10 m×10 m,20 m×20 m,25 m×25 m,50 m×50 m,100 m×100 m)的变化规律.结果显示:(1)由于多度的可加性,不同取样尺度在样地内树种多度的变化表现出一致性;随取样尺度的增加,多度方差呈线性增加,而变异系数呈线性减小.(2)丰富度随取样尺度的变化较为复杂,随取样尺度的增加,丰富度方差呈非线性变化,在取样尺度为25 m×25 m时方差最大;变异系数随取样尺度的增加而呈线性减小.研究表明,大尺度的多度值可以由小尺度的多度值通过外推法估计,而丰富度却不能,在生物多样性的保护和管理中不能简单地从一个取样尺度的生物丰富度推测另一个取样尺度丰富度.     

The role of recruitment and dispersal limitation in tree community assembly in Amazonian forests

Background: Species composition of plant communities is shaped by the interplay between dispersal limitation, environmental filters and stochastic events.

Aims: The aim of this work was to investigate the effects of dispersal limitation and environmental filtering on tree recruitment. To accomplish this, we employed the unified neutral theory of biodiversity and biogeography to examine migration within the metacommunity, defined as a set of interacting local communities linked by the dispersal of multiple potentially interacting species.

Methods: We sampled 12,975 individuals with dbh ≥ 1 cm in 26 1-ha permanent plots, including habitats of terra firme, transitional forests, várzea and campinarana, on the upper Madeira River, Brazilian Amazon.

Results: Campinarana drew individuals from outside the metacommunity species pool at a mean probability of recruitment of 0.06, a much lower probability than terra firme (0.31), transitional (0.21) and várzea forests (0.22). Environmental variables, such as water table depth, soil texture and fertility, were related to differences in community assembly.

Conclusions: Species abundance distribution and diversity patterns of plant assemblages in a large river landscape in the Amazon highlight the importance of environmental heterogeneity that conditions beta-diversity. The high variation in recruitment probabilities from the metacommunity species pool to local communities suggests high habitat variability in the process of maintaining patterns of local diversity.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Community level impacts of an ant invader and food mediated coexistence

Ant invasions exert a range of effects on recipient communities, from displacement of particular species to more complex community disruption. While species loss has been recorded for a number of invasion events, a little examined aspect of these invasions is the mechanisms for coexistence with resident ant species.The yellow crazy ant, Anoplolepis gracilipes (Smith), is considered one of the world’s worst ant invaders and has recently undergone rapid population growth in Tokelau. We surveyed the ground-dwelling ant fauna in two plots on each of five invaded and three uninvaded islands across two atolls in Tokelau to examine community characteristics of the ant fauna in areas with and without yellow crazy ants. We also used three types of food bait (tuna, jam and peanut butter) to experimentally test if species are able to coexist by consuming different food resources. Anoplolepis gracilipes was found to coexist with two to six other ant species at any one site, and coexisted with a total of 11 ant species. Four species never co-occurred with A. gracilipes. Non-metric multidimensional scaling showed significant differences in community composition and the relative abundance of species between areas that had, and had not, been invaded by A. gracilipes. The number of other ant species was significantly lower in communities invaded by the yellow crazy ant, but did not decline with increasing A. gracilipes abundance, indicating that impacts were independent of population density. The yellow crazy ant dominated all tuna and jam baits, but had a low attendance on peanut butter, allowing four other ant species to access this resource. Our results demonstrate community level impacts of an ant invader on a tropical oceanic atoll and suggest that differing use of food resources can facilitate coexistence in ant communities. Received 11 September 2006; revised 15 January 2007; accepted 22 February 2007.