Leaf litter stoichiometry affects decomposition rates and nutrient dynamics in tropical forests under restoration in Costa Rica

Active restoration strategies increase the production of leaf litter in tropical forests, but little is known about their effect on litter decomposition and subsequent nutrient release. We quantified changes in leaf litter stoichiometry during decomposition in former pasture sites under contrasting restoration strategies (natural regeneration, applied nucleation/islands tree planting and plantation), as well as in nearby primary forest. Litterbags were employed to evaluate decomposition. We used a leaf mixture of either the four planted tree species in the plantation and island treatments or the nearby primary forest and compared them under a factorial design. Decomposition rates were similar between restoration treatments (p > 0.5), but leaves decomposed faster in the forest mixture than in the plantation mixture (p < 0.01). The content of Ca, Mg, K, P, and the C:N ratio were higher in the forest mixture at the beginning and during decomposition (p < 0.05); the N content in the plantation mixture was higher at the beginning but lower during decomposition (p < 0.05), which meant greater mobilization of nitrogen per unit of carbon lost. K and P had a strong initial release, while Mg was released more gradually. N and Ca had an irregular pattern of initial fast release, immobilization, and re‐release in the later stages. We conclude that the differences in rates of decomposition and nutrient release in these systems under restoration were at least partly determined by the floristic heterogeneity and chemical quality of the leaf litter that reaches the soil.     

Planktonic primary production in a tidally influenced mangrove forest on the Pacific coast of Costa Rica

The seasonal variation of planktonic primary productivity was measured during one year in the main channel in the interior part of the mangrove forest of the Estero de Morales (Estero de Punta Morales), a mangrove system located in the Golfo de Nicoya at the Pacific coast of Costa Rica. Samples were incubated at the surface, 0.5 m and 1.0 m depth and the "light and dark bottle technique" was employed. The annual gross primary productivity (PPg) was 457 and the net primary productivity (PPn) was 278 g C m(-2) a(-1). Daily PPg ranged from 0.29 to 3.88 and PPn from 0.12 to 2.76 g C m(-2) d(-1). The highest rates observed in May and September were due to red tide blooms. The seasonal variation of primary productivity inside the mangrove forest depends closely on the PP in the adjacent area of the upper Golfo de Nicoya. Obviously the PP was light-limited since the compensation depth in the ebb current was found at only 1 m depth. In the flood current it was somewhat deeper. The planktonic primary productivity inside the mangrove forest was completely restricted to the open channels. A simultaneous measurement demonstrated that PPn of the phytoplankton could not take place under the canopy of the mangroves. Additional studies on the time course of the oxygen concentration in the mouth of the main channel over 24 hrs demonstrated a relation between the O2 and the tidal curves. The ebb current had always lower O2 concentrations than the flood current, regardless of the time of the day. The difference to the foregoing high tide, however, was much smaller when the low tide occurred during the day. This indicates that under the canopy the net primary production and hence O2 liberation of the attached macro- and microalgae, together with the high PPn of the phytoplankton in the channels, helped the oxygen concentration not to decrease as far as during the night. Nevertheless it shows that the consumtion of organic material in the submersed part of the mangrove forest exceeds always its production.     

Litter Decomposition Within the Canopy and Forest Floor of Three Tree Species in a Tropical Lowland Rain forest,Costa Rica

The rain forest canopy hosts a large percentage of the world's plant biodiversity, which is maintained, in large part, by internal nutrient cycling. This is the first study to examine the effects of site (canopy, forest floor) and tree species (Dipteryx panamensis, Lecythis ampla, Hyeronima alchorneoides) on decay rates of a common substrate and in situ leaf litter in a tropical forest in Costa Rica. Decay rates were slower for both substrates within the canopy than on the forest floor. The slower rate of mass loss of the common substrate in the canopy was due to differences in microclimate between sites. Canopy litter decay rates were negatively correlated with litter lignin:P ratios, while forest floor decay rates were negatively correlated with lignin concentrations, indicating that the control of litter decay rates in the canopy is P availability while that of the forest floor is carbon quality. The slower cycling rates within the canopy are consistent with lower foliar nutrient concentrations of epiphytes compared with forest floor-rooted plants. Litter decay rates, but not common substrate decay rates, varied among tree species. The lack of variation in common substrate decay among tree species eliminated microclimatic variation as a possible cause for differences in litter decay and points to variation in litter quality, nutrient availability and decomposer community of tree species as the causal factors. The host tree contribution to canopy nutrient cycling via litter quality and inputs may influence the quality and quantity of canopy soil resources.     

Vegetation effects on microclimate in lowland tropical forest in Costa Rica

The temperature and atmospheric humidity in a tropical lowland rain forest in Costa Rica were measured in order to assess the microclimate in different forest environments. Two disturbed sites, a single tree fall gap (400 m²) and an 0.5 hectare clearing, were compared for periods up to two years after disturbance. Two locations in primary forest, the canopy and understory, were also monitored. Temperatures were highest in the clearing, intermediate in the canopy and gap which were similar, and lowest in the understory. Vapor pressure deficits (VPD) were highest in the clearing, followed by the canopy, the gap and the understory. With regrowth of the vegetation in the gap and clearing sites, the temperatures and vapor pressure deficits significantly decreased. After 1 year, the microclimate at seedling height in the clearing resembled that of the gap, and after two years the microclimate of the gap was very similar to that of the understory. Seasonal differences in temperature and VPD were small compared to differences caused by changes in the stature of the vegetation.     

The Landscape Ecology of Tropical Secondary Forest in Montane Costa Rica

Multinomial logistic models of land use/land cover in montane Costa Rica and landscape pattern analysis showed that relative to agriculture, secondary forest occurred closer to old-growth forest, further from roads, in forest reserves, and at higher elevations. Collinearity between explanatory variables yielded simple multivariate models; proportion of surrounding old growth predicted secondary forest most accurately. An old-growth matrix [mean patch size (MPS) 24.5 ha], located mainly within protected areas, dominated elevations greater than 2500 m. A matrix of agriculture (MPS 23.5 ha), with smaller patches (approximately 9 ha) of secondary forest and old growth, dominated elevations from 1500 to 2500 m. Combining secondary forest with old growth decreased forest patch number and increased MPS from 7.3 to 37.1 ha. I concluded that: (a) secondary forest pattern is nonrandom, so ancillary data will aid its mapping with satellite imagery. The variables elevation, agriculture distance, road distance, and population density distinguished secondary forest from old growth with 74% accuracy; (b) socioeconomic and biological forces probably interact to create these secondary forest patterns; and (c) the strong association between secondary forest and old growth supports the concept that tropical forest recovery depends on the landscape structure of remnant forest. Received 16 February 1999; accepted 20 August 1999.     

Tropical Montane Forest Restoration in Costa Rica: Overcoming Barriers to Dispersal and Establishment

Tropical forests are being cleared at an alarming rate although our understanding of their ecology is limited. It is therefore essential to design restoration experiments that both further our basic knowledge of tropical ecology and inform management strategies to facilitate recovery of these ecosystems. Here we synthesize the results of research on tropical montane forest recovery in abandoned pasture in Costa Rica to address the following questions: (1) What factors limit tropical forest recovery in abandoned pasture? and (2) How can we use this information to design strategies to facilitate ecosystem recovery? Our results indicate that a number of factors impede tropical forest recovery in abandoned pasture land. The most important barriers are lack of dispersal of forest seeds and seedling competition with pasture grasses. High seed predation, low seed germination, lack of nutrients, high light intensity, and rabbit herbivory also affect recovery. Successful strategies to facilitate recovery in abandoned pastures must simultaneously overcome numerous obstacles. Our research shows that establishment of woody species, either native tree seedlings or early‐successional shrubs, can be successful in facilitating recovery, by enhancing seed dispersal and shading out pasture grasses. On the contrary, bird perching structures alone are not an effective strategy, because they only serve to enhance seed dispersal but do not reduce grass cover. Remnant pasture trees can serve as foci of natural recovery and may enhance growth of planted seedlings. Our results highlight the importance of: (1) understanding the basic biology of an ecosystem to design effective restoration strategies; (2) comparing results across a range of sites to determine which restoration strategies are most generally useful; and (3) considering where best to allocate efforts in large‐scale restoration projects.     

The effect of litter type and macrofauna community on litter decomposition and organic matter accumulation in post-mining sites

Field microcosms consisting of mineral soil (spoil substrate) and two types of litter taken either from an unreclaimed site with spontaneously developed vegetation (mostly Salix caprea) or from an alder plantation (a mixture of Alnus glutinosa and A. incana) were exposed in spontaneously developed or reclaimed sites at a post-mining heap near Sokolov (Czech Republic) for one year. The litter types differed remarkably in C:N ratio which was 29 for spontaneous litter and 14 for alder litter. The two microcosm types were either accessible or not accessible to soil macrofauna. The effect of macrofauna exclusion on soil mixing was complex and depended on litter quality and the site that determined soil fauna composition. In reclaimed sites where macrofauna was dominated by saprophags, mainly earthworms, the macrofauna access increased soil mixing. In sites where predators dominated, the macrofauna exclusion probably suppressed fragmentation and mixing activity of the mesofauna.     

Snail populations,beech litter production,and the role of snails in litter decomposition

Summary The population densities of snails living in beech litter were studied form March 1968 to April 1969. Litter production over one year was measured and the role of snails in litter disappearance assessed.Snails were extracted from litter using a modified Vágvölgyi (1952) flotation method, extraction efficiencies being 84%. The mean annual population density of the twenty-one species of snail recorded on the main sampling site was estimated at 489/m². Carychium tridentatum was the most numerous species, with a mean density of 200/m². Acanthinula aculeata, Punctum pygmaeum and Vitrea contracta also had fairly high mean densities. The mean annual biomass was 699 mg dry wt./m² or 278 mg ash-free dry wt./m². Hygromia striolata and Oxychilus cellarius/alliarius were the most important species in terms of biomass on the main site. Within the limits of accuracy imposed by the sampling regime the population densities of four out of five of the species (C. tridentatum, A. aculeata, V. contracta, Retinella pura) studied remained unchanged throughout the year, whereas P. pygmaeum had a significantly higher autumn population. C. tridentatum populations were highly aggregated at all times of the year, most markedly so in June. Other species were aggregated at certain times of the year only. Samples taken from other sites showed total population densities of snails ranging from 185–1082 snails/m².A total tree litter production of 652 g/m²/annum was recorded of which 584g/m²/annum was of beech material. 72% fell in the October–December period. 58% of the beech litter-fall was leaves, 5.2% bud-scales, 27% fruits and 10% twigs and bark. Summation of appropriate field layer peak standing crops amounted to 23.3 g/m². This was considered as potential litter and was equivalent to 3.4% of the total litter input. The litter standing on the woodland floor in Septermber 1968 was 2,700 g/m², hence, assuming a steady state, litter turnover time was estimated as 4.5 years.It was calculated that the total snail population ingested 0.35–0.43% of the annual litter input, of which 49% was assimilated. The role of the individual species is examined in relation to concepts of key species in ecosystem functioning. The possible role of slugs in decomposition processes is also discussed.     

Establishment of Epiphytic Bromeliads in Successional Tropical Premontane Forests in Costa Rica

Plant community composition is the combined result of species-specific competitive abilities and the availability of propagules. For epiphytic plants, current hypotheses consider that dispersal-related factors are most important. By controlling seed dispersal constraints, we experimentally examined whether the community composition of epiphytic bromeliads in a tropical premontane area is determined during early phases of seedling recruitment. Also, we tested whether establishment success was related to eco-physiological traits of the species. A total of 7200 seeds were artificially affixed on several host trees in two secondary forest patches and in a mature forest stand. Four bromeliad species with differing physiological characteristics (CAM, C₃-CAM, and C₃) and habitat preference (secondary vs. primary forest) were selected. We found that differences in seed germination probability among habitats and species were not likely to influence community assembly. After 2 yr, seedling survival and plant development were relatively higher in the early-successional forest. Seedling establishment success was not associated with specific physiological and morphological adaptations or habitat preference of the studied species. Our results were not consistent with the described community composition and rates of population recruitment of the studied species in the same successional habitats. The results support the hypothesis that chance and historic events related to seed dispersal have an important influence on community assembly of epiphytic plants. In addition, differences in growth rates and reproductive turnover among species are expected to influence the relative abundance and recruitment rates in a particular habitat.     

Microhabitat distribution of protostelids in a Tropical Wet Forest in Costa Rica

A microhabitat study of protostelids was carried out in a Tropical Wet Forest at the La Selva Biological Station in Costa Rica. Nine species were recorded from sterile wheat straws placed out and then re-collected over a period of six weeks from two different litter microhabitats in an area of primary forest. All nine species were present on straws placed in the aerial litter microhabitat, but only six species were present on straws placed in the forest floor litter microhabitat. Total colonies, percent of straws colonized, and mean number of species per straw increased significantly over time. One species (Schizoplasmodiopsis pseudoendospora) typical of temperate litter was the overwhelming dominant on the forest floor litter, while Echinostelium bisporum, a species rare in temperate litter microhabitats, was the single most abundant species in the aerial litter microhabitat. Both of these species had significantly increased frequencies over time. Two species abundant in temperate aerial litter microhabitats and one species abundant in temperate forest floor litter were rare at La Selva. Our data conform to those obtained in an earlier study carried out in tropical forests in the mountains of Puerto Rico and provide additional support towards developing a model of microhabitat distribution of protostelids in terrestrial ecosystems.     

森林凋落物的微生物分解

森林凋落物的分解是森林生态系统中物质循环和能量流动的一个重要环节,而微生物在这一过程中起着重要作用。本文系统介绍了森林凋落物微生物分解的过程及其生态学意义,并从参与凋落物分解的微生物多样性、凋落物分解过程中的微生物数量动态及群落演替、影响微生物分解的因素及微生物分解酶学等方面综述了森林凋落物的微生物分解研究概况,探讨了未来研究的方向。     

The effect of climate and soil conditions on tree species turnover in a Tropical Montane Cloud Forest in Costa Rica

On a global level, Tropical Montane Cloud Forests constitute important centers of vascular plant diversity. Tree species turnover along environmental gradients plays an important role in larger scale diversity patterns in tropical mountains. This study aims to estimate the magnitude of beta diversity across the Tilardn mountain range in North-Western Costa Rica, and to elucidate the impact of climate and soil conditions on tree species turnover at a local scale. Seven climate stations measuring rainfall, horizontal precipitation (clouds and wind-driven rain) and temperatures were installed along a 2.5km transect ranging from 1200 m.a.s.l. on the Atlantic to 1200 m.a.s.l. on the Pacific slope. The ridge top climate station was located at 1500 m.a.s.l. Climate data were recorded from March through December 2003. Additionally, seven 0.05 ha plots were established. On all plots soil moisture was monitored for one year, furthermore soil type and soil chemistry were assessed. Woody plants with a diameter at breast height (dbh) > or = 5 cm were identified to species. Species' distributions were explored by feeding pairwise Serensen measures between plots into a Principal Component Analysis. Relationships between floristic similarity and environmental variables were analyzed using Mantel tests. Pronounced gradients in horizontal precipitation, temperatures and soil conditions were found across the transect. In total, 483 woody plants were identified, belonging to 132 species. Environmental gradients were paralleled by tree species turnover; the plots could be divided in three distinctive floristic units which reflected different topographic positions on the transect (lower slopes, mid slopes and ridge). Most notably there was a complete species turnover between the ridge and the lower Pacific slope. Floristic similarity was negatively correlated with differences in elevation, horizontal precipitation, temperatures and soil conditions between plots. It is suggested that beta-diversity in the study area is largely driven by species with narrow spatial ranges, due to the interactions between topography, climate and soil formation processes, especially around the wind-exposed and cloud covered ridge area. The findings emphasize the extraordinary conservation value of tropical montane cloud forests in environmentally heterogeneous areas at mid-elevations.     

Effect of litter substrate quality and soil nutrients on forest litter decomposition: A review

Forest litter plays an important role in determining nutrient cycling, balance and maintaining ecosystem function of forest ecosystems. Studies have shown that litter substrate quality is one of the most important factors affecting litter decomposition in a given area. It is, hence, important to understand the factors controlling litter decomposition in the late decomposition stage and determining organic matter changes over the duration of litter decomposition. Decomposition rate of mixed litter may differ with that of a single specie litter. Supply of soil nutrients is an important factor controlling litter decomposition rate, because the essential nutrients in soil or litter material influence community and activity of decomposers (soil organisms). There were clear relationships among soil nutrient, litter substrate quality, and decomposition. Soil nutrient contents were positively correlated with litter substrate quality, showing that higher contents of soil nutrient were accompanied with good quality of litter substrate, and lower soil nutrients with poor litter quality. The effects of soil fertility on litter decomposition rate varied with environmental conditions. It was reported that litter quality regulates the early stage of carbon decomposition and its accumulation in soil, however, it could not predict the long-term dynamics of soil organic carbon. Hence, the formation and stabilization of soil organic carbon are controlled by the quantity of litter input and its interaction with the soil circumstances rather than by the litter quality. The present paper reviewed the research findings about litter decomposition related to litter substrate quality and soil nutrients, including short-term and long-term litter decomposition, decomposition of single-species vs. mixed-litter decomposition and litter nutrients release. The present paper aimed to clarify the relationship between soil nutrients and litter decomposition, which will help to understand forest succession, forest water conservation and soil re-production capacity.     

The occurrence of Frankia in tropical forest soils of Costa Rica

Infective and effective Frankia were shown to occur in five diverse tropical forest soils of Costa Rica. Results of a plant infection assay indicated that Frankia is a common component of the soil biota in low and high elevation, primary and secondary forest soils. This is the first report of Frankia in lowland tropical rainforests of the Americas. These results suggest either a nonsymbiotic population of soil Frankia, the presence of unknown actinorhizal host species, or an ability of Frankia to be dispersed over long distances.     

氮沉降对森林凋落物分解的影响

氮沉降增加作为全球变化的重要现象之一,已经并将继续对森林凋落物分解产生影响.综述了国内外氮沉降对森林凋落物分解影响及其机理的研究现状.氮沉降对凋落物分解的影响可分为直接影响和间接影响.氮沉降通过影响森林地被物组成和凋落物化学成分,间接影响凋落物分解.氮沉降对凋落物分解的直接影响表现为促进、无影响和抑制3种效果.分析了产生以上影响效果的作用机理,介绍了氮沉降对森林凋落物分解影响的研究方法,探讨了目前研究存在的问题,讨论了未来该方面研究的重点和方向.     

Effects of freeze damage on litter production, quality and decomposition in a loblolly pine forest in central China

Background and aims

Freeze events can strongly influence many ecosystem processes. However, the effects of freeze events on litter production, litter quality, and decomposition are rarely documented.

Methods

In this study, litter fall was measured monthly for 2 years. Two litter decomposition experiments were also performed using freeze-damaged litter and non-damaged litter in a loblolly pine forest.

Results

The freeze event in November 2009 caused a pronounced pulse of needle litter fall. The freeze-damaged needle litter was shown to have higher N concentration and lower C/N ratio compared with the normal falling needle litter. This finding indicates that freeze damage significantly increased needle quality because of incomplete nutrient resorption. The decomposition of freeze-damaged needle litter was faster than that of normal falling yellow needle litter and slower than that of hand-picked green needle litter. The decomposition rate constant (k) was negatively correlated with the C/N ratio in the needle litter. Our results also showed that the different climatic conditions influence patterns of litter decomposition.

Conclusions

This study suggests that freeze events significantly alter litter quantity and quality, thus affecting litter decomposition rates in a loblolly pine forest in central China.     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Nesting bird species in sun coffee, pasture, and understory forest in southern Costa Rica

We investigated avian nest distribution and success in understoryforest, sun coffee plantations, and pasture in southern Costa Rica. Nestsearching occurred in plantations and forest in 1999 and 2000 and in pastures in2000. Nests were monitored until they failed or fledged young. Antbirds(Thamnophilidae) were the most common understory forest nesters and were notfound nesting in the plantations or pastures. Common nesting species in theplantations included Turdidae, Tyrannidae, Cardinalidae, and Thraupidae, many ofwhich are typical of forest edge/canopy or open, scrubby habitats. Two speciesassociated with forest interior, Henicorhina leucostictaand Buarremon brunneinucha, were found nesting in theplantations. Pastures supported similar types of nesting species as theplantations, with the exception of the forest-interior species. Daily mortalityrates (DMRs) for above-ground cup-nesting species in plantations and pastureswere similar to those for species nesting in forest at our site and a site inPanama. The results indicate that conversion from forest to pastures and suncoffee plantations diminishes nesting habitat for forest-interior species, whilenumerous forest edge/canopy species and open-country species are able to nest inthese agricultural land-cover types. As a group, species nesting in theplantations and pastures do not have unusually high nest mortality rates,although species-specific studies are lacking. Nesting species distributionsacross habitat types and DMRs at our study site may be influenced by the largeamount of forest in the landscape.