Variation in leaf litter nutrients of a Costa Rican rain forest is related to precipitation

By assessing current leaf litter nutrient dynamics, we may be able to predict responses of nutrient cycling in tropical ecosystems to future environmental change. The goal of this study was to assess whether nutrient cycling is related to seasonal variation in rainfall in a wet tropical forest. We examined leaf litter of an old-growth tropical rain forest in N.E. Costa Rica over a 4-year period to explore seasonal and inter-annual changes in leaf litter nutrient concentrations, and to evaluate potential short- and long-term drivers of variation in litter nutrient concentration, particularly that of phosphorus (P) and nitrogen (N). We also examined the temporal dynamics of calcium, potassium, and magnesium in the leaf litter. Leaf litter [P] and %N changed significantly with time, both seasonally and inter-annually. Seasonal changes in leaf litter [P] were strongly positively correlated with rainfall from the previous 2 weeks; cations, however, were inversely related to this measure of current rainfall, while %N was not related to rainfall. We propose that the positive relationship between current rainfall and leaf litter [P] is due to a response by the vegetation to an increase in nutrient availability and uptake. In contrast, given the negative relationship between current rainfall and cation concentrations, leaching from live leaf tissue is a more likely driver of short-term changes in cations. Should global climate change include altered rainfall patterns in this biome, one class of ecosystem-level responses could be significant changes in P and cation cycling.     

Photosynthesis and successional status of Costa Rican rain forest trees

Seven tree species from three different light environments in the wet lowland forests of Costa Rica were grown under controlled environment conditions to assess light related photosynthetic potentials. Light saturated photosynthesis rates were clearly related to light levels of the field environments. Mean saturated, net photosynthetic rates ranged from 6.8 to 11.3 to 27.7 mol m^–2 sec^–1 for plants from heavy shade, canopy light gaps and man-made clearings respectively. Light saturation of plants from clearings occurred at photosynthetic photon flux densities greater than 1000 mol m^–2 sec^–1 whereas plants from heavy shade environments became light saturated near 500 mol m^–2 sec^–1. Plants that normally occur in intermediate light environments were intermediate in light saturation levels. Mean maximum stomatal conductances ranged from 1.0 to 7.3 mm sec^–1 and followed a pattern similar to photosynthetic rates.     

Removal of seeds from frugivore defecations by ants in a Costa Rican rain forest

At our Costa Rican field site, seeds defecated by frugivorous birds usually do not remain where they have been deposited. Many species of ants are attracted to frugivore defecations and remove seeds and/or pulp. Pheidole species selectively remove seeds, fungus-growing species (tribe Attini) remove both pulp and seeds. Seeds of many Melastomataceae have an appendage, which we hypothesized is an elaiosome. Indeed, preference trials demonstrated that two species of Pheidole selected seeds with the appendage over seeds of the same species in which the appendage had been removed. However, we found that these ants did not take the appendage when it was offered by itself. We conclude that the appendage is not an elaiosome. In further trials, different ant species preferentially selected different seed species. These ants consumed some seeds and deposited others unharmed in refuse piles. We conclude that because the composition of leaf-litter ant communities is highly variable between neighboring square meter plots, and the probability of seed predation depends upon the species of ant, the over-all effect of ants on seed shadows and seed banks is spatially unpredictable. Addendum: The names of the two Pheidole emphasized in this study. P. nebulosa and P. nigricula, are unpublished names from a generic revision being prepared by E. O. Wilson and W. L. Brown. Their use here is not intended to constitute taxonomic publication but is solely for more precise indentification in future ecological research of similar nature     

Microclimatic parameters influencing nitrogen fixation in the phyllosphere in a Costa Rican premontane rain forest

The acetylene reduction method was used to measure nitrogen fixation in the phyllosphere of attached leaves of different phorophytes under natural conditions in a premontane rain forest in Costa Rica. Maximum rates of nitrogen fixation (26 ng N · cm⁻² leaf area · h⁻¹) – mainly due to the activity of two species of Scytonema (Cyanobacteria) – were measured in the rainy season in bright sunlight. Rates of nitrogen fixation were correlated with the leaf area covered by Scytonema. In periods without precipitation the fixation activity decreased to zero within 2–3 days. As long as the epiphylls were sufficiently supplied with water, other microclimatic factors like temperature and light intensity also influenced nitrogen fixation rates, but to a lesser extent. Relative humidity and species of phorophyte showed no direct influence. It was concluded that the most important factor for nitrogen fixation in the phyllosphere was the availability of liquid water. Linking these results to meteorological data, the input of nitrogen by biological nitrogen fixation in the phyllosphere in the investigation area was estimated to be as much as 1.6 ± 0.8 kg N · ha⁻¹ · year⁻¹ per unit of leaf area index (LAI). For an LAI of 2 for the understory the nitrogen input would vary between 2 and 5 kg N · ha⁻¹ · year⁻¹. This work also demonstrates the importance of detailed knowledge of variation in microclimate throughout the year as a basis for extrapolation of the annual nitrogen input. Received: 21 December 1997 / Accepted: 14 June 1998     

Root surface acid phosphatase activities of vascular epiphytes of a Costa Rican rain forest

The objective of this work was to compare root surface phosphatase activities of vascular epiphytes typical of a lowland tropical forest. Acid phophatase, measured at pH 5.0 with the substrate p-nitrophenyl phosphate, was detected in 22 species distributed within 10 plant families. Epiphytes were classified as trunk, canopy-mat or bare-limb species based upon their usual occurrence. Phosphatase activity was not significantly correlated with plant occurrence. However, phosphatase activity was generally highest in trunk occurring and canopy-mat epiphytes rooted in mosses and humus-like accumulations, and lowest in species restricted to bare limbs. Epiphyte shoot phosphorus and chlorophyll content were correlated with species occurrence, with phosphatase being positively correlated with plant P content. The observed changes in acid phosphatase production among habitats were consistent with predicted changes in the availability of organic P sources. However, observed changes also reflect accompanying shifts in root structure including: the occurrence of velamen or waxy layers, changes in root diameter, branching and root hair density.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Swimming ability in three Costa Rican dry forest rodents

We investigated the swimming abilities of three Costa Rican dry forest rodents (Coues' rice rat. Oryzomys couesi, hispid cotton rat, Sigmodon hispidus, and spiny pocket mouse, Liomys salvini) associated with a large marsh, Laguna Palo Verde, using 90 s swim trials in a plastic container. Swimming ability was evaluated by observing the use of limbs and tail in the water, inclination to the surface, and diving and floating behavior. Rice rats could float, swim and dive, suggesting that they can exploit surface and underwater resources. Cotton rats swam at the water's surface, but were less skilled swimmers than rice rats. Spiny pocket mice tired quickly and had difficulty staying at the water's surface. Results suggest that differential swimming ability is related to the distribution of the three sympatric species within the marsh and adjacent forest habitats.     

Substrate conditions,foliar nutrients and the distributions of two canopy tree species in a Costa Rican secondary rain forest

A 28 yr old secondary lowland rain forest in Costa Rica was dominated by two tree species of contrasting ecologies, Vochysia ferruginea - a species typical of well-drained but infertile soils of high Al saturation, and Cordia alliodora, a species requiring soils of at least moderate fertility. The two species exhibited markedly different spatial distributions in the forest and we sought to determine whether or not these different distributions were related to variation in substrate conditions and the nutrient requirements of the two species. Two soil types were present: Ultisols (Typic Haplohumults) and Inceptisols (Typic Dystropepts) and topography was of low hills. The study formed part of an analysis of site quality for timber production in the two species and therefore used standard forestry methods for such analyses. 36 plots of 20 m×20 m were established using predetermined criteria of site uniformity, presence of at least four dominant trees of at least one of the study species, and absence of disturbance. The abundance (number of individuals 10 cm dbh) of each species was determined in each plot. Soil samples were taken in 27 of the 36 plots, 9 plots being considered to replicate conditions already sampled, and 14 soil chemical and physical variables were measured using standard methods. Foliar nutrient analyses were carried out for trees of both species with crowns fully exposed to sunlight, distributed across the gradient of soil conditions. Variations in species abundances were not related to soil type, but plots in which Vochysia ferruginea was more abundant tended to be on steeper slopes with soils of higher exchangeable acidity and lower concentrations of Mn; Cordia alliodora was more abundant on gentler topography where soils had much lower exchangeable acidity but higher Mn. Differences between the two species in foliar nutrient concentrations were marked and supported previous interpretations of their nutrient requirements; foliar nutrients of Vochysia ferruginea were typical of tree species of moist tropical forest on infertile soils, and those of Cordia alliodora typical of a species requiring more fertile soils. Factors such as the distributions of seed trees at site abandonment may affect the spatial distributions of tree species in secondary forests such as that studied. The relationships of the distributions of the two species to substrate variation, however, and their evidently different nutrient requirements, support the hypothesis that variation in the composition and structure of the forest studied is at least partially related to exchangeable acidity and its dominant cation, Al.     

White-lipped peccary (Tayassu pecari,Artiodactyla: Tayassuidae) diet and fruit availability in a Costa Rican rain forest

We studied fruit availability, diet and habitat use by white-lipped peccary (Tayassu pecari) in Corcovado National Park, southwest Costa Rica, from July 1996 to April 1997. The results show that the availability of important fruits for the white-lipped peccary differs between habitats and climatic seasons. Fruit availability was highest in the primary forest than secondary and coastal forest. There was a period of shortage of fruits to ends of the wet season, during which the consumption of not seasonal resources like leaves and shafts increased. The important fruits during this period of shortage were Ficus sp and Licania operculipetala. The several types of forest were used according to the fruit availability, and it was a direct relation between the consumption and the fruit availability.     

Epiphytic and terrestrial mycorrhizas in a lower montane Costa Rican cloud forest

The epiphyte community is the most diverse plant community in neotropical cloud forests and its collective biomass can exceed that of the terrestrial shrubs and herbs. However, little is known about the role of mycorrhizas in this community. We assessed the mycorrhizal status of epiphytic (Araceae, Clusiaceae, Ericaceae, and Piperaceae) and terrestrial (Clusiaceae, Ericaceae) plants in a lower montane cloud forest in Costa Rica. Arbuscular mycorrhizas were observed in taxa from Araceae and Clusiaceae; ericoid mycorrhizas were observed in ericaceous plants. This is the first report of intracellular hyphal coils characteristic of ericoid mycorrhizas in roots of Cavendishia melastomoides, Disterigma humboldtii, and Gaultheria erecta. Ericaceous roots were also covered by an intermittent hyphal mantle that penetrated between epidermal cells. Mantles, observed uniquely on ericaceous roots, were more abundant on terrestrial than on epiphytic roots. Mantle abundance was negatively correlated with gravimetric soil water content for epiphytic samples. Dark septate endophytic (DSE) fungi colonized roots of all four families. For the common epiphyte D. humboldtii, DSE structures were most abundant on samples collected from exposed microsites in the canopy. The presence of mycorrhizas in all epiphytes except Peperomia sp. suggests that inoculum levels and environmental conditions in the canopy of tropical cloud forests are generally conducive to the formation of mycorrhizas. These may impact nutrient and water dynamics in arboreal ecosystems.     

冻融环境下凋落叶添加对亚高山森林土壤腐殖化程度的影响

亚高山森林凋落叶与土壤腐殖化的关系是了解森林土壤有机质动态过程的关键内容,且受到季节性冻融、凋落叶与土壤性质的综合作用,但缺乏深入认识。因此,在对亚高山凋落叶腐殖化认识的前期基础上,以川西亚高山典型的针叶林、针阔混交林以及阔叶林土壤为研究对象,采用控制冻融环境和凋落叶添加的方法,通过分析土壤可提取腐殖质的色调系数(Δlog K)、光密度值(E4/E6)以及A600/C值等光学指标,研究冻融循环与凋落叶添加对土壤腐殖化程度的影响。结果表明:冻融循环和凋落叶添加及其交互作用均明显影响土壤的腐殖化程度,但受到凋落叶类型和土壤类型的调控。在冻融循环初期,添加凋落叶能促进土壤的腐殖化进程,提高土壤的腐殖化程度;随着冻融循环次数的增加以及凋落叶的不断分解,阔叶凋落叶对土壤的腐殖化表现出促进作用,而针叶凋落叶则对土壤的腐殖化表现出抑制作用。此外,在冻融循环环境下,即使土壤的腐殖化程度增加,但腐殖质仍为相对年轻的Rp型,有利于提高土壤的肥力。这些结果表明,气候变化情景下高山森林地表冻融循环的改变将显著影响凋落叶与土壤腐殖质的密切关系,为进一步认识区域森林土壤有机质动态提供了一定的科学依据。     

Sex ratio and breeding of white-lipped peccaries Tayassu pecari (Artiodactyla: Tayassuidae) in a Costa Rican rain forest.

White-lipped peccaries are non-seasonal breeders in South America, but little is known about their reproduction in Central America. There are few studies about the sex ratio of this species in the field. We studied the reproduction and sex ratio of white-lipped peccaries during 200 hours of field observation of four radiomarked and two unmarked herds, from July 1996 to April 1997, in Corcovado National Park, Costa Rica. Sex ratio data of three additional, radiomarked herds observed in 1998 were also included. We recorded numbers of mountings, presence of newborns and numbers of nursing interactions. The peccaries showed a distinct reproductive seasonality, with one mating peak in February and another in July. The greatest number of newborns and the peak in nursing activity were observed during July and August, when fruit availability for the peccaries was high. The adult sex ratio was significantly female biased (1.4:1-1.8:1), also in contrast with South American populations.     

Evidence for dispersal of fig seeds by the fruit-eating characid fish Brycon guatemalensis Regan in a Costa Rican tropical rain forest

Studies were conducted at the La Selva Biological Station in the Atlantic lowlands of Costa Rica and in a greenhouse in California to assess the potential role of a Neotropical fish in dispersing the seeds of a rain forest tree. Feeding experiments showed that the seeds of Ficus glabrata H. B. K., a major, canopy-forming riparian tree, require approximately 18–36 h to pass through the digestive tract of Brycon guatemalensis Regan, an abundant riverine fish whose adult diet consists largely of leaves and fruits of this fig tree. The seeds were still viable after passing through the fish’s gut but germinated somewhat more slowly than seeds that had been left in the fig exposed to air or floated in water. Stem elongation of seedlings from seeds that had passed through the fish’s gut was faster than that of seeds in the other two treatments. Placement of seeds upstream may be more important than enhanced germination for plants such as Ficus that produce large numbers of seeds. Radio telemetry showed that five of six tagged fish had moved distances of 0.1–1 km upstream; seven other fish with transmitters, including three large males, were not relocated and may have moved into tributary streams for spawning or feeding. These findings suggest that Brycon can disperse large numbers of Ficus seeds and help maintain the upstream populations of the tree. Received: 1 May 1996 /Accepted: 1 August 1996     

Influence of land crabs Gecarcinus quadratus (Gecarcinidae) on distributions of organic carbon and roots in a Costa Rican rain forest

In Costa Rica's Corcovado National Park, the fossorial land crab, Gecarcinus quadratus (Gecarcinidae), densely populates (1 - 6 m(-2)) a region of forest extending from the Park's Pacific coastline inland to ca. 600 m. Throughout this coastal forest ('crabzone'), crabs selectively forage for fallen leaves and relocate them to subterranean burrow chambers. Comparisons between surface soils (0 - 15 cm) sampled from the crabzone and forest lying immediately inland that is naturally devoid of crabs ('crabless zone') suggest that crabzone top soils contained less organic carbon and fewer fine and very fine roots. In contrast, soils sampled from 70 - 100 cm depths in the crabzone contained twice the carbon of the crabless zone during the dry season but similar values during the wet season. Two years of experimental crab exclusion from 25 m2 replicates established in the crabzone resulted in 16% more organic carbon content in surface soils relative to baseline conditions (n.s.) and 22% more carbon than final control values (P < 0.05). Excavations of burrow chambers, with an average (+/- SD) depth of 48 +/- 12 cm, indicated localized, subterranean pockets of elevated (+ 60 %) organic carbon and increased densities of fine and very fine roots relative to same-depth samples from the crabzone unassociated with burrows chambers.     

Rodent pollination of Blakea (Melastomataceae) in a Costa Rican cloud forest

Several species of rodents have been observed visiting the flowers of an undescribed species ofBlakea (Melastomataceae) in a cloud forest in Costa Rica. The rodents drink the copious nectar secreted at the base of the stamens and are dusted with pollen. The first report of pollination by non-flying mammals in the neotropics suggests that this type of pollination may be important where there is little competition with bats.     

Carbon allocation in a Costa Rican dry forest derived from tree ring analysis

The rising discussion on carbon balance of tropical forests often does not consider the sequestration potential of secondary dry forests, which are becoming an increasing importance due to land use change and reforestation. We have developed an easy applicable tool for the estimation of biomass increment of tropical secondary forest stands on the base of tree ring analysis. The existence of annual rings was shown by a combination of anatomical examination and radiocarbon estimations. With tree ring analysis, forest inventories and destructive sampling the above-ground biomass increment of secondary forest stands of age between 9 and 48 years in the dry forest region of Guanacaste, Costa Rica were estimated. The above-ground biomass increment of the tree layer varies between 2.4 and 3.2 Mg/ha yr in different stands. Lianas contribute with up to 23% additional production. Differences in productivity among the stands along a chronosequence were not significant. The measured carbon allocation potential of 1.7–2.1 Mg C/ha yr lies in the range of reported values from other tropical dry forests and old growth humid forests as well.     

Changes in diversity along a successional gradient in a Costa Rican upper montane Quercus forest

Changes in terrestrial vascular plant diversity along a successional gradient were studied in a Costa Rican upper montane Quercus forest. In 1991 and 1992 species presence and cover were recorded in 12 successional 0.1 ha forest plots. A total of 176 species in 122 genera and 75 families were found. Asteraceae was the most speciose family. With the help of TWINSPAN three successional phases were classified: (i) Early Secondary Forest (ESF, 145 spp.), (ii) Late Secondary Forest (LSF, 130 spp.) and (iii) Primary Forest (PF, 96 spp.). Detrended Correspondence Analysis (DCA) species ordination using DECORANA illustrates that different ecological species groups can be distinguished along the time sequence. Alpha diversity (Shannon-Wiener index, among others) in ESF and LSF was significantly greater than in PF. This is probably explained by downslope migration of numerous sub(alpine) species to cleared and recently abandoned montane sites. Beta diversity applying Sørensen's similarity coefficients declined during succession. Using linear regression, the minimum time required for floristic recovery following disturbance and abandonment was calculated at 65.9 years. A comparison with other studies shows that secondary forests in upper montane Costa Rica can be as diverse as in neotropical lowlands.     

山地森林表层土壤酶活性对短期增温及凋落物分解的响应

为了探究贡嘎山地区暗针叶林表层土壤酶活性对增温及凋落物分解的响应,采用开顶式生长箱(open top chamber)和加热电缆(OTC-cable)联合增温的方式模拟增温,同时定位监测实验小区地表空气、表层土壤温湿度的变化;不同类型(A:75%峨眉冷杉针叶+25%杜鹃荚蒾灌木叶凋落物,B:55%峨眉冷杉针叶+45%杜鹃荚蒾灌木叶凋落物)凋落物在模拟增温和自然对照条件下分解4年,研究土壤过氧化氢酶、多酚氧化酶和脲酶活性的变化。结果表明:增温使得地表空气和表层土壤温度分别增加了2.84℃和1.83℃;使得空气相对湿度和土壤含水量分别降低了5.27%和1.55%。针叶比例高会抑制凋落物分解,约13%;增温促进凋落物分解且对针叶比例高的促进作用更加明显,增幅均超过10%。增温总体上降低了土壤过氧化氢酶和脲酶活性,而对多酚氧化酶活性的影响表现为增大。针叶比例降低的凋落物分解对3种土壤酶活性的影响大致趋势是增大,幅度在15%以上。增温和凋落物类型之间的交互作用显著。随着土壤深度增加,土壤过氧化氢酶和多酚氧化酶活性增大,而脲酶活性降低。增温和不同类型凋落物分解对表层土壤酸碱性无显著作用。在土壤含水量较低的情况下,土壤水分对酶活性影响较大。贡嘎山峨眉冷杉林表层土壤酶对温度的敏感性不仅因酶类型、土壤深度而存在差异,也随增温时间、土壤水分条件而有所不同。     

Asymmetric effects of litter removal and litter addition on the structure and function of soil microbial communities in a managed pine forest

Aims

The effect of different MeJA doses applied prior to or simultaneously with toxic Al on biochemical and physiological properties of Vaccinium corymbosum cultivars with contrasting Al resistance was studied.

Methods

Legacy (Al-resistant) and Bluegold (Al-sensitive) plants were treated with and without toxic Al under controlled conditions: a) without Al and MeJA, b) 100 μM Al, c) 100 μM Al + 5 μM MeJA, d) 100 μM Al + 10 μM MeJA and e) 100 μM Al + 50 μM MeJA. MeJA was applied to leaves 24 h prior to or simultaneously with Al in nutrient solution. After 48 h, Al-concentration, lipid peroxidation (LP), H₂O₂, antioxidant activity, total phenols, total flavonoids, phenolic compounds and superoxide dismutase activity (SOD) of plant organs were analyzed.

Results

Al-concentrations increased with Al-treatment in both cultivars, being Al, LP and H₂O₂ concentrations reduced with low simultaneous MeJA application. Higher MeJA doses induced more oxidative damage than the lowest. Legacy increased mainly non-enzymatic compounds, whereas Bluegold increased SOD activity to counteract Al³⁺.

Conclusions

Low MeJA doses applied simultaneously with Al³⁺ increased Al-resistance in Legacy by increasing phenolic compounds, while Bluegold reduced oxidative damage through increment of SOD activity, suggesting a diminution of its Al-sensitivity. Higher MeJA doses could be potentially toxic. Studies are needed to determine the molecular mechanisms involved in the protective MeJA effect against Al-toxicity.

     

Fine-root biomass and soil properties in a semideciduous and a lower montane rain forest in Panama

The distribution of root biomass and physical and chemical properties of the soils were studied in a semideciduous and in a lower montane rain forest in Panama. Roots and soil samples were taken by means of soil cores (25 cm deep) and divided into five, 5-cm deep sections. Soils were wet-sieved to retrieve the roots that were classified in four diameter classes: very fine roots (<1 mm), fine roots (1–2 mm), medium roots (2–5 mm) and coarse roots (5–50 mm). Soil samples were analyzed for organic carbon, total nitrogen, available phosphorus, exchangeable bases, cation exchange capacity, pH, aluminium and exchangeable acidity. Total root biomass measured with the soil corer (roots <50 mm in diameter) was not different between the forests (9.45 t ha^-1), while biomass of very fine roots was larger in the mountains (2.00 t ha^-1) than in the lowlands (1.44 t ha^-1). The soils in the semideciduous forest were low in available phosphorus, while in the mountains, soils had low pH, high exchangeable aluminium and exchangeable acidity, and low concentration of exchangeable bases. Phosphorus was in high concentration only in the first 5 cm of the soil. In both forests, there was an exponential reduction of root biomass with increasing depth, and most of the variation in the vertical distribution of roots less than 2 mm in diameter was explained by the concentration of nitrogen in the soils. The results of this study support the hypothesis that a large root biomass in montane forests is related to nutrients in low concentration and diluted in organic soils with high CEC and low bulk density, and that fine root biomass in tropical forests in inversely related to calcium availability but not a phosphorus as has been suggested for other forests.