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.     

Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems

The widespread use of forest litter as animal bedding in central Europe for many centuries gave rise to the first litter manipulation studies, and their results demonstrated that litter and its decomposition are a vital part of ecosystem function. Litter plays two major roles in forest ecosystems: firstly, litterfall is an inherent part of nutrient and carbon cycling, and secondly, litter forms a protective layer on the soil surface that also regulates microclimatic conditions. By reviewing 152 years of litter manipulation experiments, I show that the effects of manipulating litter stem from changes in one, or both, of these two functions, and interactions between the variables influenced by the accumulation of litter can result in feedback mechanisms that may intensify treatment effects or mask responses, making the interpretation of results difficult.Long-term litter removal increased soil bulk density, overland flow, erosion, and temperature fluctuations and upset the soil water balance, causing lower soil water content during dry periods. Soil pH increased or decreased in response to manipulation treatments depending on forest type and initial soil pH, but it is unclear why there was no uniform response. Long-term litter harvesting severely depleted the forests of nutrients. Decreases in the concentrations of available P, Ca, Mg, and K in the soil occurred after only three to five years. The decline in soil N occurred over longer periods of time, and the relative loss was greater in soils with high initial nitrogen concentration. Tree growth declined with long-term litter removal, probably due to lower nutrient availability. Litter manipulation also added or removed large amounts of carbon thereby affecting microbial communities and altering soil respiration rates.Litter manipulation experiments have shown that litter cover acts as a physical barrier to the shoot emergence of small-seeded species; further, the microclimate maintained by the litter layer may be favourable to herbivores and pathogens and is important in determining later seedling survival and performance. Litter manipulation altered the competitive outcomes between tree seedlings and forbs, thereby influencing species composition and diversity; changes in the species composition of understorey vegetation following treatments occurred fairly rapidly. By decreasing substrate availability and altering the microclimate, litter removal changed fungal species composition and diversity and led to a decline in populations of soil fauna. However, litter addition did not provoke a corresponding increase in the abundance or diversity of fungi or soil fauna.Large-scale long-term studies are still needed in order to investigate the interactions between the many variables affected by litter, especially in tropical and boreal forests, which have received little attention. Litter manipulation treatments present an opportunity to assess the effects of increasing primary production in forest ecosystems; specific research aims include assessing the effects of changes in litter inputs on the carbon and nutrient cycles, decomposition processes, and the turnover of organic matter.     

A test of the hypothesis that correlational selection generates genetic correlations

Theory predicts that correlational selection on two traits will cause the major axis of the bivariate G matrix to orient itself in the same direction as the correlational selection gradient. Two testable predictions follow from this: for a given pair of traits, (1) the sign of correlational selection gradient should be the same as that of the genetic correlation, and (2) the correlational selection gradient should be positively correlated with the value of the genetic correlation. We test this hypothesis with a meta-analysis utilizing empirical estimates of correlational selection gradients and measures of the correlation between the two focal traits. Our results are consistent with both predictions and hence support the underlying hypothesis that correlational selection generates a genetic correlation between the two traits and hence orients the bivariate G matrix.     

Multiple hypothesis testing to detect lineages under positive selection that affects only a few sites

Detection of positive Darwinian selection has become ever more important with the rapid growth of genomic data sets. Recent branch-site models of codon substitution account for variation of selective pressure over branches on the tree and across sites in the sequence and provide a means to detect short episodes of molecular adaptation affecting just a few sites. In likelihood ratio tests based on such models, the branches to be tested for positive selection have to be specified a priori. In the absence of a biological hypothesis to designate so-called foreground branches, one may test many branches, but a correction for multiple testing becomes necessary. In this paper, we employ computer simulation to evaluate the performance of 6 multiple test correction procedures when the branch-site models are used to test every branch on the phylogeny for positive selection. Four of the methods control the familywise error rates (FWERs), whereas the other 2 control the false discovery rate (FDR). We found that all correction procedures achieved acceptable FWER except for extremely divergent sequences and serious model violations, when the test may become unreliable. The power of the test to detect positive selection is influenced by the strength of selection and the sequence divergence, with the highest power observed at intermediate divergences. The 4 correction procedures that control the FWER had similar power. We recommend Rom's procedure for its slightly higher power, but the simple Bonferroni correction is useable as well. The 2 correction procedures that control the FDR had slightly more power and also higher FWER. We demonstrate the multiple test procedures by analyzing gene sequences from the extracellular domain of the cluster of differentiation 2 (CD2) gene from 10 mammalian species. Both our simulation and real data analysis suggest that the multiple test procedures are useful when multiple branches have to be tested on the same data set.     

Using a botanic garden collection to test a bioclimatic hypothesis

Bioclimatic hypotheses are used to explain how climate regulates the occurrence of species. A derivative of these hypotheses is that plants moved between corresponding bioclimatic areas should thrive, whereas plants moved to a different zone should languish. This principle is routinely applied in forestry and horticulture but actual tests of the hypotheses seem scanty. We carried out a test on the Finnish system of bioclimatic vegetation zoning using the plant collection of Helsinki University Botanic Garden in Kumpula, which is situated at the northern limit of the hemiboreal zone. We aimed to test how the plants’ survival depends on their provenance with the expectation that plants from the hemiboreal or southern boreal zones should do best in Kumpula. Probability of survival was estimated using collection database information of 379 plant accessions of known wild origin, and logit models. Different growth forms were analysed separately. In most analyses accessions of temperate and hemiarctic origin showed lower survival probability than those originating from any of the boreal subzones, which among them exhibited rather evenly high probabilities. Trees were an exception showing an almost steadily increasing survival probability from temperate to northern boreal origin. In all, the results gave some support to the tested hypothesis, but the various factors that could not be controlled for produced results that were difficult to interpret. We conclude that botanic gardens should pay due attention to information management and curational practices to ensure widest possible applicability of their plant collections.     

Differential palatability of leaf litter to four sympatric isopods in a Hong Kong forest

Summary The feeding behaviour of four sympatric isopods from a Hong Kong forest has been investigated. The study included two armadillids (Formosillo raffaelei and Orodillo maculatus) and two philosciids (Burmoniscus ocellatus and Burmoniscus sp.). When given a choice of eight types of litter, all isopod species showed significant dietary selection, and food preferences were similar. Berchemia racemosa (Rhamnaceae) was most readily consumed, followed by Celtis sinensis (Ulmaceac), while feeding rates on Cinnamomum camphora (Lauraceae) were low. The armadillids exhibited a narrower dietary spectrum than the philosciids. All isopods showed significant differences in consumption rates when each litter type was presented separately to them. Cinnamomum camphora was eaten the slowest, while Celtis sinensis and Berchemia racemosa were the top-ranked species. Although patterns of litter ranking based on percentage eaten or feeding rates were similar, all isopods ate more food when given a mixture of leaves than when presented with a single litter type. Assimilation rates were, in general, positively related to feeding rates. On this basis, it appears that assimilation may influence food preference. There were no clear relationships between food preference or feeding rates and ash, calcium, copper, soluble tannin or energy content of the litter. This finding may indicate the benefits of isopods maintaining a mixed diet, consuming certain litter species to meet their calcium or copper requirements and then switching to others so as to meet daily energy needs and to avoid excessive injection of tannins or plant allelochemicals.     

Edge and herbivory effects on leaf litter decomposability in a subtropical dry forest

It is increasingly recognized that understanding the functional consequences of landscape change requires knowledge of aboveground and belowground processes and their interactions. For this reason, we provide novel information addressing insect herbivory and edge effects on litter quality and decomposition in fragmented subtropical dry forests in central Argentina. Using litter from Croton lachnostachyus (a common shrub species in the region) in a decomposition bed experiment, we evaluated whether litter quality (carbon and nitrogen content; carbon: nitrogen ratio) and decomposability (percentage of remaining dry weight) differ between litter from forest interiors or edges (origin) and with or without herbivory (damaged/undamaged leaves). We found that edge/interior origin had a strong effect on leaf litter quality (mainly on carbon content), while herbivory was associated with a smaller increase in nitrogen content. Herbivore damage increased leaf litter decomposability, but this effect was related to origin during the initial period of litter incubation. Overall, undamaged leaf litter from the forest edge showed the lowest decomposability, whereas damaged leaf litter decomposed at rates similar to those observed in litter from the forest interior. The interacting edge and herbivory effects on leaf litter quality and decomposability shown in our study are important because of the increasing dominance of forest edges in human-modified landscapes and the profound effect of leaf litter decomposition on nutrient cycling.     

Involvement of lignocellulolytic enzymes in the decomposition of leaf litter in a subtropical forest

The involvement of ligninolytic and cellulolytic enzymes, such as laccase, lignin peroxidase, manganese peroxidase, carboxymethylcellulase (CMCase), and filter paper activity (FPA), in the decomposition process of leaf litter driven by 6 soil-inhabiting fungi imperfecti was studied under solid-state fermentations. All the tested fungi exhibited varied production profiles of lignocellulolytic enzymes and each caused different losses in total organic matter (TOM) during decomposition. Based on the results, the 6 fungi could be divided into 2 functional groups: Group 1 includes Alternaria sp., Penicillium sp., Acremonium sp., and Trichoderma sp., and Group 2 includes Pestalotiopsis sp. and Aspergillus fumigatus. Group 1, with higher CMCase and FPA activities, showed a higher decomposition rate than the fungi of Group 2 over the first 16 d, and thereafter the cellulolytic activities and decomposition rate slowed down. Group 2 showed the maximum and significantly higher CMCase and FPA activities than those of the Group 1 fungi during the later days. This, combined with the much higher laccase activity, produced a synergistic reaction that led to a much faster average mass loss rate. These results suggest that the fungi of Group 1 are efficient decomposers of cellulose and that the fungi of Group 2 are efficient decomposers of lignocellulose. During cultivation, Pestalotiopsis sp. produced an appreciable amount of laccase activity (0.56+/-0.09 U/ml) without the addition of inducers and caused a loss in TOM of 38.2%+/-3.0%, suggesting that it has high potential to be a new efficient laccase-producing fungus.     

亚热带不同树种凋落叶分解对氮添加的响应

为探究不同质量凋落物对氮(N)沉降的响应, 该研究采用尼龙网袋分解法, 在亚热带福建三明格氏栲(Castanopsis kawakamii)自然保护区的米槠(Castanopsis carlesii)天然林, 选取4种本区常见的具有不同初始化学性质的树种凋落叶进行模拟N沉降(N添加)分解实验(施N水平为对照0和50 kg·hm ^-2·a ^-1)。研究结果表明: 在2年的分解期内, 对照处理的各树种凋落叶的分解速率依次为观光木(Michelia odora, 0.557 a ^-1)、米槠(0.440 a ^-1)、台湾相思(Acacia confusa, 0.357 a ^-1)、杉木(Cunninghamia lanceolata, 0.354 a ^-1); N添加处理凋落叶分解速率依次为观光木(0.447 a ^-1)、米槠(0.354 a ^-1)、杉木(0.291 a ^-1)、台湾相思(0.230 a ^-1), 除杉木凋落叶外, N添加显著降低了其他3种凋落叶分解速率。N添加不仅使4种树木凋落叶分解过程中的N释放减慢, 同时还抑制凋落叶化学组成中木质素和纤维素的降解; N添加在凋落叶分解过程中总体上提高β-葡萄糖苷酶(βG)和酸性磷酸酶活性, 对纤维素水解酶的活性影响不一致, 而降低β-N-乙酰氨基葡萄糖苷酶活性和酚氧化酶活性。凋落叶分解速率与凋落叶中的碳获取酶(βG)活性以及其化学组分中的可萃取物含量极显著正相关, 与初始碳浓度、纤维素和木质素含量极显著负相关, 与初始N含量没有显著相关性。凋落物类型和N添加的交互作用虽未影响干质量损失速率, 但对木质素和纤维素的降解具有显著效应。综上所述, 化学组分比初始N含量能更好地预测凋落叶分解速率, 而N添加主要通过抑制分解木质素的氧化酶(如PHO)来降低凋落叶分解速率。     

利用凋落物法和林木因子模拟小兴安岭阔叶红松林叶面积指数

叶面积指数(LAI)是研究森林生态系统生理生态进程中关键的结构参数之一。目前,凋落物法是在非破坏性条件下能直接测定森林生态系统LAI的最有效的方法,然而将凋落叶按树种分类增加了该方法的实施难度。平均优势度模型、林分优势度模型和局域优势度模型基于凋落物法和林木因子(如胸高断面积basal area,BA;坐标)能精确地预测落叶阔叶林的LAI,而这些模型是否适用于针阔混交林仍未进行验证。以小兴安岭阔叶红松林(Pinus koraiensis)为研究对象,先利用凋落物法测定其LAI,依此为参考对3种模型预测LAI的有效性进行验证,并以红松、冷杉(Abies nephrolepis)、紫椴(Tilia amurensis)、五角槭(Acer mono)、枫桦(Betula costata)和裂叶榆(Ulmus laciniata)为例,探讨了基于凋落物法测定的LAI与BA的相关关系。结果表明:平均优势度模型不适于预测针阔混交林的LAI;林分优势度模型预测效果较好,精度达86%;局域优势度模型预测效果最优,精度高于90%。然而,为准确测定阔叶红松林的LAI,应最少选择测定8个主要树种的比叶面积。基于凋落物法测定的6个树种的LAI与其BA均显著相关(P0.01),最小R2为0.67。研究结果可为快速、准确地测定针阔混交林的LAI提供依据,为非破坏性条件下建立树种的LAI与其BA的相关关系提供参考。     

Microfungi associated with Abies needles and Betula leaf litter in a subalpine coniferous forest

We investigated microfungal assemblages on leaf litter within a subalpine forest in central Japan and their variation with season, litter depth, and litter species. Microfungal assemblages were compared for Abies needles and Betula leaf litter collected from litter and fermentation layers of the forest floor during the growing season in spring, summer, and autumn. A total of 35 and 42 species were isolated from Abies needles and Betula leaf litter, respectively. The observed variation in microfungal assemblages was primarily attributable to seasonal differences. The frequencies of Trichoderma viride, Volutella ciliata, Mucor sp., and Umbelopsis ramanniana increased in summer, leading to a high degree of similarity of microfungal assemblages in different litter depths and litter species. Microfungal assemblages on Abies needles in spring and autumn and those on Betula leaves in spring were characterized by Trichoderma viride, V. ciliata, Thysanophora penicillioides, Trichoderma polysporum, and (or) Mortierella alpina. Microfungal assemblages on Betula leaves in autumn were characterized by the absence of these species and the occurrence of Cladosporium cladosporioides. The results were discussed with an emphasis on the role of microfungi in decomposition processes and the impact on fungi of predicted future increases in global temperature.     

Diversity and host recurrence of fungi associated with the bleached leaf litter in a subtropical forest

Fungi that selectively remove lignin cause extensive bleaching of leaf litter, which results in the acceleration of litter decomposition. The taxonomic diversity and host recurrence of saprotrophic fungi associated with bleached leaf litter were investigated in a subtropical forest in southern Japan. A total of 211 fungal isolates and sporocarps were obtained from bleached portions of leaf litter of 33 plant species in 18 families and were classified into 83 operational taxonomic units (OTUs) at the 97% similarity level of the ITS rDNA sequence. These fungal OTUs belonged to Rhytismataceae and Xylariaceae in the Ascomycota, and Marasmiaceae, Mycenaceae, Lachnocladiaceae, and Physalacriaceae in the Basidiomycota. OTUs in Rhytismataceae and Marasmiaceae showed a significantly higher degree of recurrence for plant species than simulated networks where partners were associated randomly. In contrast, OTUs in Xylariaceae and Mycenaceae showed no significant recurrence for plant species. Such differing degrees of recurrence for plant species implied different roles of fungal families in leaf litter decomposition.     

Using model selection to choose a size-based condition index that is consistent with operational welfare indicators

Quantitative and qualitative measures of fish health and welfare are essential for management of both wild capture and aquaculture species. These measures include morphometric body condition indices, energetic condition and aquaculture operational welfare indicators (OWIs). Measures vary in ease of measurement (and may require destructive sampling), and it is critical to know how well they correlate with fish health and welfare so appropriate management decisions can be based on them. Lumpfish (Cyclopterus lumpus) is a new farming species that needs nondestructive OWIs to be developed and validated. In this study, we developed a C. lumpus fin damage score. Four different body condition indexes based on individual weight relative to either length–weight relationships or relative to other fish in its local environment were tested (using model selection) as predictors of individual fin damage. Results showed severity of fin damage was predicted by small size relative to the other individuals in the tank or cage. Body condition based on length–weight relationship was not found to predict fin damage, indicating that using established indices from fisheries or from other species would not predict welfare risks from fin damage. Implications are that especially in hatchery conditions grading will improve the condition index, and is expected to mitigate fin damage, but that low weight at length was not of use in predicting fin damage. Model selection to choose between a suite of possible indices proved powerful and should be considered in other applications where an easily measured index is needed to correlate with other health measures.     

Human skin-color sexual dimorphism: a test of the sexual selection hypothesis

Applied to skin color, the sexual selection hypothesis proposes that male preference for light-skinned females explains the presence of light skin in areas of low solar radiation. According to this proposal, in areas of high solar radiation, natural selection for dark skin overrides the universal preference of males for light females. But in areas in which natural selection ceases to act, sexual selection becomes more important, and causes human populations to become light-skinned, and females to be lighter than males. The sexual selection hypothesis proposes that human sexual dimorphism of skin color should be positively correlated with distance from the equator. We tested the prediction that sexual dimorphism should increase with increasing latitude, using adult-only data sets derived from measurements with standard reflectance spectrophotometric devices. Our analysis failed to support the prediction of a positive correlation between increasing distance from the equator and increased sexual dimorphism. We found no evidence in support of the sexual selection hypothesis.     

Resource utilization and guild structure of small vertebrates in the Amazon forest leaf litter

Ten species of small vertebrates that occurred together in leaf litter of the Amazon tropical rainforest were studied during the dry season in Rondônia, western Brazil. This vertebrate assemblage consisted of two diurnal lizards, two diurnal frogs, one nocturnal frog, and five frog species that are active by day and night. All species feed on small invertebrates, primarily insects and arachnids. There are significant differences in prey types and sizes eaten, and overlap values among species are relatively low. Niche breadths vary from unity (extremely low) to high (> 9.0). A pseudocommunity analysis using two different algorithms indicated that structure exists in the prey resource matrix of the vertebrate assemblage. The zero structure of the community matrix contributes to the observed structure, i.e. prey items not taken by certain species are important in maintaining structure. Several well-defined feeding guilds are apparent, and the feeding guilds do not appear to follow taxonomic lines.     

Biodegradation ofShorea robusta Gaertn. leaf litter and the cycling of minerals in a tropical sal forest

Summary Microbial degradation of the leaf litter ofShorea robusta Gaertn. was studied for a period of one year. The changes in the major litter constituents like sugar, starch, hemicellulose, cellulose and lignin were analysed from the litter kept under the nylon bags, at a month's interval. Four major elements phosphorous, potassium, magnesium and calcium were also assessed monthly from the decaying litter and the soil. The role of microfungi in mineral cycling and loss of litter substrates have been correlated.     

Environmental rugosity, body size and access to food: a test of the size-grain hypothesis in tropical litter ants

In terrestrial walking organisms, long legs help to decrease the cost of running, allowing animals to step over environmental interstices rather than walking through them. However, long legs can complicate the infiltration of these interstices, which may contain food sources and refugia. Since the number of environmental interstices perceived by an organism (rugosity) increases as it body size decreases (size-grain hypothesis, SGH), natural selection should favor proportionally smaller legs with decreasing body size. Recent work demonstrated that ants fit this hypothesis. We experimentally tested the assumption of the SGH that small ants, which have proportionally smaller legs than larger ants, are more successful in exploring environmental interstices because they can easily penetrate them. We examined the ability of tropical litter ant species with different body sizes to access food baits in 'landscapes' (=plots) with different levels of rugosity and food exposure. In the first experiment, three levels of landscape rugosity were defined by manipulating the density of leaf litter placed on the ground plots: a) plain landscape: no litter fall, b) intermediate rugosity (∼0.5 kg of litter fall) and c) high rugosity (∼1 kg). In a second experiment, food baits were in plain landscapes, exposed or covered by leaf litter. The body lengths of ants that first accessed food baits ranged from 1.5 to 12 mm. Ants that first reached food baits in the most rugose landscapes were ∼40% smaller than ants that first found baits in plain landscapes. Smaller ants were also the first to access covered food. The application of a phylogenetic comparative method suggested the same patterns. We conclude that these results support the size grain hypothesis. Environmental rugosity might have operated as a selective force to shape the morphological characteristics of litter ant species.     

The interaction between developmental bias and natural selection: from centipede segments to a general hypothesis

Do limitations to the ways in which mutations can alter developmental processes help to determine the direction of phenotypic evolution? In the early days of neo-Darwinism, the answer given to this question was an emphatic 'no'. However, recent work, both theoretical and empirical, argues that the answer should at least be 'sometimes', and possibly even a straightforward 'yes'. Here, I examine the key concept of developmental bias, which encompasses both developmental constraint and developmental drive. I review the case of centipede segment number, which is a particularly clear example of developmental bias, but also a rather unusual one. I then consider how, in general terms, developmental bias and natural selection might interact, with the result that it is their interaction, rather than either process on its own, that determines evolutionary direction. Essentially, the whole argument is about the extent to which phenotypic variation is developmentally structured as opposed to amorphous or random. This issue can be traced back to the very beginning of evolutionary biology, and in particular to a difference of opinion between Darwin and Wallace, who emphasized, respectively, character correlation and character independence.     

Temporal shifts in the relative importance of climate and leaf litter traits in driving litter decomposition dynamics in a Chinese transitional mixed forest

Plant and Soil - To assess the direction and strength of climate and leaf litter trait effects on decomposition dynamics throughout the litter decomposition process. We performed a three-year-long...