Determinants of Lichen Diversity in a Rain Forest Understory

Change in lichen diversity is often used as a bioindicator to estimate effects of atmospheric pollution, but natural variation in lichen cover and species richness can be very high. We examined the top-down effects of spore-consuming ants and the bottom-up effects of nutrient and light availability on lichen diversity associated with the leaf surface of the rain forest understory plant, Piper cenocladum. Plots containing P. cenocladum were randomly assigned to treatments in factorial experiments that included high and low light levels, nutrient enrichment, and presence and absence of the ant mutualist, Pheidole bicornis . At the conclusion of the experiments, plants were harvested and size of leaves, secondary metabolite content (amides), epiphyll cover, and the species richness of the lichens (which comprised 85% of the epiphyll community) were quantified. Epiphyll cover (mosses, liverworts, and lichens) was greater on plants that had ant-mutualists and balanced resources. Lichen species richness was greater for plants with balanced resources, particularly for those with high light availability. Relationships between toxins and lichen cover and richness were weak and unclear. In this system, natural sources of variation were reliable determinants of lichen diversity and both biotic and abiotic influences were important.     

Impact of Atta colombica Colonies on Understory Vegetation and Light Availability in a Neotropical Forest1

We quantified patterns of vegetation removal and light availability above Atta colombica nests on Barro Colorado Island, Panama. Ants cleared vegetation less than 1 cm in diameter from an area of 77 m², and up to 3 m above ground level. Overall light availability 1.5 m above ground level was 49 percent greater at ant nest sites than at sites in undisturbed understory. These higher light levels fell within the range known to enhance growth of both shade tolerant and pioneer species.     

Influence of Microclimate on the Occurrence of Cyanobacteria in the Phyllosphere in a Premontane Rain Forest of Costa Rica

Abstract: The occurrence of cyanobacteria in the phyllosphere of several understory plants was investigated during a one and a half year study in a primary premontane rain forest in Costa Rica. Altogether, seven species of epiphyllous cyanobacteria were identified and are discussed here. The ecology of the two most frequent species in the understory, Scytonema javanicum and Sc. hofmannii , was studied in further detail. Within the Understory, the relative abundance of these species, as well as of epiphyllous bryophytes, was more influenced by air humidity than by light. The humidity, characterized as potential evaporation, also had influence on the pseudosuccession of the phyllosphere vegetation. While at moist sites bryophytes and both species of Scytonema appeared nearly simultaneously on 6–9-month-old leaves, at dryer sites Sc. javanicum , and especially Sc. hofmannii , appeared 6–9 months after bryophytes had established. On 13–15-month-old leaves bryophytes and Scytoriema were usually well established. On average, 20–30% of the leaf area of 2–5-year old leaves of Spathacanthus hoffmannii (Acan-thaceae, an abundant understory shrub) were covered by bryophytes, 2–3% by Sc. javanicum , and 0.1–0.2% by Sc. hofmannhi . An influence of phorophyte species on the occurrence and abundance of these two species of Scytonema could not be detected. However, Sc. hofmannii was more frequent on leaves with higher bryophyte cover, while Sc. javanicum was independent from bryophytes. A comparison of leaves from the understory and from the canopy showed that the composition and abundance of cyanobacteria species changed with height above forest floor.     

Lack of Interspecific Plant Competition With a Dominant Grass in the Understory of a Lowland Forest in Colombia

I analyzed a natural experiment of competition involving a dominant herb in Tinigua Park, Colombia. Pharus virescens is a dominant monocarpic grass that synchronously produces seeds once every 5 yr, after which all individuals die. I found no difference in species richness, density, or diversity comparing plots with and without the grass in 381 quadrats (4 m²) and there was no negative relationship between the abundance of the grass and that of other plants.     

The Importance of Where to Dump the Refuse: Seed Banks and Fine Roots in Nests of the Leaf‐Cutting Ants Atta cephalotes and A. colombica1

The location of the nutrient‐rich organic refuse produced by a leaf‐cutting ant colony varies among ant species. Atta cephalotes locate their organic refuse in subterranean chambers, whereas A. colombica place their organic refuse on the soil surface near the nest. We studied the effect of the absence or presence of external organic refuse on the abundance of fine roots and seed bank composition in the superficial horizons of ant nests. We sampled soils from ant nests or dumps and adjacent areas of 15 adult nests of A. cephalotes at La Selva (LS), Costa Rica, and of 15 of A. colombica nests on Barro Colorado Island (BCI), Panama. Soils from A. cephalotes nests did not differ from adjacent soils in abundance of fine‐root and seed diversity. In contrast, organic refuse from A. colombica nests was less diverse in seed composition (due to the great abundance of Miconia argentea) and had a greater abundance of fine roots than adjacent areas. Thus the external location of the ant‐nest organic refuse is potentially important in determining the different types of plant recolonization in abandoned or dead ant nests. The relative abundance of these Atta species may influence the structure and/or composition of tropical forests.     

Effects of Dispersal Pattern and Mammalian Herbivores on Seedling Recruitment for Virola michelii (Myristicaceae) in French Guiana1

The effects of dispersal pattern (seeds in small clumps vs. seeds scattered in pairs) and distance to the nearest Carapa procera (Meliaceae; a tree that produces seeds preferred by terrestrial vertebrates) on survival of seeds and seedlings were examined for the animal–dispersed tree species Virola michelii (Myristicaceae) in a mature forest‘at Paracou, French Guiana, in 1992 and 1993. We assessed the putative role of ground–dwelling mammalian herbivores, rodents, and ungulates that filter the seed shadow, acting either as dispersers or predators and thus modifying the original pattern of seed dispersal made by frugivores. We measured the effects of simulated seed burial by rodents using marked seeds and quantified the effect of protecting seeds and seedlings from ground–dwelling vertebrates on seedling germination and survival with fence exclosures in 1992. Dispersal pattern had short–term but no long–term effects on the proportion of V. michelii seeds that survived one year later as seedlings. In the short term, within six weeks, clumped seeds survived better than scattered seeds in both years. Marked seeds that were removed from their site of dispersal were eaten; rodents only rarely buried seeds of V. michelii, and seed burial reduced seed and seedling survivorship. The combined effect of the factors year and Carapa proximity significantly affected seed survival within six weeks. Although six–week seed survival was greater in 1993 than in 1992, seedling establishment was lower in 1993 than in 1992 following a lower rainfall regime during the key period of seed germination (February). One–year seed and seedling survivorship was similar between treatments and years. Seed survival and seedling establishment in V. michelii was dependent on vertebrates in the short term and on climate in the long term. Overall, seed and seedling survivorship depended on a combination of these factors.     

Edge Effects of Linear Canopy Openings on Tropical Rain Forest Understory Microclimate

We investigated microclimatic edge gradients associated with grassy powerlines, paved highways and perennial creeks in wet tropical forest in northeastern Australia during wet and dry seasons. Photosynthetically active radiation, air temperature and vapor pressure deficit, soil temperature, canopy temperature, soil moisture, and air speed in the rain forest understory were measured during traverses perpendicular to the forest edge. Light intensity was elevated near the edges of powerlines, highways, and creeks, but this effect was strongest for creek edges. Air temperature and vapor pressure deficit were elevated near powerline edges in the dry season and highway edges in both wet and dry seasons but were not elevated near creek edges in either season. In contrast, soil moisture was lowered near creek edges but not near either powerline or highway edges. No edge gradients were detected for air speed. Canopy temperature was elevated near highway edges and lowered near powerline edges in the wet season but no edge gradients in canopy temperature were detected near creek edges in either the wet or the dry season. We suggest that these different edge gradients may be largely the result of differences in the fluxes of latent and sensible heat within each type of linear canopy opening, with periodic flood disturbance assisting by maintaining a more open canopy near creek edges. Our data indicate that the nature of the linear canopy opening is at least as important as the width in determining the nature and severity of microclimatic edge effects, analogous to the "matrix effect" of traditional fragmentation studies.     

Ecology of Foliicolous Lichens at the “Botarrama” Trail (Costa Rica), a Neotropical Rain Forest. I. Species Composition and its Ecogeographical Implications1

A total of 217 species of typically foliicolous lichens were found at the “Botarrama” trail, a premontane rain forest in the Braulio Carrillo National Park, Costa Rica. The species composition confirmed the classification of this study sice as transitional between lowland wet and montane rain forest. In terms of species tichness, the foliicolous lichen flora was dominated by the families Gomphillaceae (Calenia, Echinoplaca, Gyalideopsis, and Tricharia), Trichotheliaceae (Porina and Tricbothelium), and Pilocarpaceae (Byssoloma and Fellhanera). The most frequent species were Porina mirabilis (including its supposed anamorph Phyllophiale alba), Gyalectidium filicinum, Porina rufula, and Strigula platypoda. A large proportion of the species was rare. Frequency distributions based on area cover corresponded well to general community models by closely fitting a log-normal approximation. Half of the species had narrow ecological amplitudes while more than 70 percent exhibited a wide geographical distribution, thus making che study site representative of a Neotropical rain forest. The use of foliicolous lichens as indicators of altitudinal zonation is discussed.     

Heterogeneity of Light Availability and its Effects on Simulated Carbon Gain of Tree Leaves in a Small Gap and the Understory in a Tropical Rain Forest1

To clarify the small-scale heterogeneity of light regimes in a rain forest, photosynthetic photon flux density (PFD) was measured at 1-min intervals during six days at 12 microsites in each of two plots, a small gap and an understory in Pasoh Forest Reserve, Peninsular Malaysia. Frequency distribution of microsite PFD was unimodal with the peak value between 16 and 32 μmol/m²/sec in the small gap, but between 8 and 16 μmol/m²/sec in the understory. In the small gap, PFD was more variable among microsites; total daily PFD and daily sunfleck PFD exceeding 10 μmol/ m²/sec tended to be higher (P <0.05; t-test) compared to those in the understory. Sunfleck PFD exceeding 50 μmol/ m²/sec, however, showed no difference between the two plots. Diffuse PFD transmittance, defined as the ratio of PFD in the forest to that measured at 43 m above ground during the periods 0800-0810 and 1750-1800 h, was significantly higher in the small gap than in the understory plot. Diffuse PFD transmittance was also positively correlated with microsite total daily PFD. To examine the effects of the subtle heterogeneity of light regimes on leaf carbon gain, we simulated carbon gain by sun and shade leaves in a typical shade-tolerant species, Brosimum aticastrum Sw. (Moraceae). Despite the similarity in total daily PFD, total daily carbon gain was considerably higher in the gap than in the understory for both sun and shade leaves. This study suggests that frequency distribution of PFD is critical in describing microsite PFD regimes and determining leaf carbon gain in the tropical forest floor.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Effects of Inbreeding,Outbreeding, and Supplemental Pollen on the Reproduction of a Hummingbird‐pollinated Clonal Amazonian Herb

Understory herbs are an essential part of tropical rain forests, but little is known about factors limiting their reproduction. Many of these herbs are clonal, patchily distributed, and produce large floral displays of nectar‐rich 1‐d flowers to attract hummingbird pollinators that may transport pollen over long distances. The aim of this study was to investigate the effects of clonality, cross‐proximity, and patchy distribution on the reproduction of the hummingbird‐pollinated Amazonian herb Heliconia metallica. We experimentally pollinated flowers within populations with self‐pollen and with pollen of different diversity, crossed flowers between populations, and added supplemental pollen to ramets growing solitarily or in conspecific patches. Only flowers pollinated early in the morning produced seeds. Selfed flowers produced seeds, but seed number and mass were strongly reduced, suggesting partial sterility and inbreeding depression after selfing. Because of pollen competition, flowers produced more seeds after crosses with several than with single donor plants. Crosses between populations mostly resulted in lower seed production than those within populations, suggesting outbreeding depression. Ramets in patches produced fewer seeds than solitary ramets and were more pollen‐limited, possibly due to geitonogamy and biparental inbreeding in patches. We conclude that high rates of geitonogamy due to clonality and pollen limitation due to the short receptivity of flowers and patchy distribution constrain the reproduction of this clonal herb. Even in unfragmented rain forests with highly mobile pollinators, outbreeding depression may be a widespread phenomenon in plant reproduction.     

Effects of experimental nitrogen additions on plant diversity in an old‐growth tropical forest

Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N‐limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old‐growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N‐mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha^?1 yr^?1 (Low‐N); 100 kg N ha^?1 yr^?1 (Medium‐N), and 150 kg N ha^?1 yr^?1 (High‐N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low‐to‐medium levels of N addition (≤100 kg N ha^?1 yr^?1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine‐root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition‐based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high‐N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.     

Effects of Canopy Gaps,Topography, and Soils on the Distribution of Woody Species in a Central Brazilian Deciduous Dry Forest1

The interrelationships between the distribution of woody species and environmental variables were investigated in an area of deciduous dry forest in Santa Vitöria, central Brazil. This is the first study of a vanishing type of dry forest which grows on base-rich soils originating from the basalt bedrocks of southern Goiás and western Minas Gerais. A survey of topography, soil properties, canopy gaps and woody plants (≥5 cm diameter at the base of the stem) was conducted in 50–15 × 15 m quadrats. The soils were classified into the following soil series: Hapludolls → Haplustolls → Haplustolls → Ustropepts → Rhodustalfs. This series corresponded to a gradient of increasing elevation and effective soil depth and decreasing slope gradient, soil organic matter and total exchangeable bases. A canonical correspondence analysis and a detrended correspondence analysis indicated that plant species’abundance distribution was significantly correlated with both the relative area of canopy gaps in the quadrats and the soil-topography gradient. Presumably, the critical factors involved in these two gradients are, respectively, light and ground water regimes. The influence of canopy gaps (i.e., light) was surprising and has not been documented previously for tropical deciduous dry forests.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Effects of Logging on the Diversity of Lianas in a Lowland Tropical Rain Forest in Hainan Island, South China

Lianas are an integral part of tropical forest ecosystems, which usually respond strongly to severe disturbances, such as logging. To compare the effect of different logging systems on the lianas diversity in tropical rain forest, we recorded all lianas and trees ≥1 cm dbh in two 40-year-old forest sites after clear cutting (CC) and selective cutting (SC) as well as in an old-growth (OG) lowland tropical rain forest on Hainan Island in south China. Results showed that OG contained fewer liana stems and lower species richness (stems: 261, richness: 42 in 1 ha) than CC (606, 52) and SC (727, 50). However, OG had the highest Fisher's α diversity index (17.3) and species richness per stem (0.184). Species composition and dbh class distribution of lianas varied significantly with different logging systems. The mean liana dbh in OG (22.1 cm) were higher than those in CC (7.0 cm) and SC (10.4 cm). Stem twining was the most frequent climbing mechanism represented in the forest, as shown by the greatest species richness, abundance, basal area, and host tree number with this mechanism. The percent of host tree stems ≥4 cm dbh hosting at least one liana individual in SC (39%) was higher than CC (23%) and OG (19.5%). Large host trees (dbh≥60 cm) were more likely to be infested by lianas in SC and OG. Our study demonstrated that logging disturbance could significantly change the composition and structure of liana communities in the lowland tropical rain forest of south China.
     

Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.