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In recent years, three sections (Humipedon, Copedon and Lithopedon) were recognized in the soil profile. It was then possible to link the first and most biologically active section to the characteristics of the environment and soil genesis. In particular, it is now possible to distinguish organic horizons, mainly produced by arthropods and enchytraeids in cold and acidic or dry and arid environments, from organo-mineral horizons produced by earthworms in more temperate and mesotrophic environments. Each set of horizons can be associated with a humus system or form, with important implications for forestry. Anecic/endogeic earthworms and Mull or Amphi systems are more abundant in the early and late stages of sylvogenesis; by completely recycling litter, earthworms accelerate the availability of organic and inorganic soil nutrients to roots and pedofauna. On the other hand, arthropods and Moder or Tangel systems characterize the intermediate stages of sylvogenesis, where thickening in the organic horizons and the parallel impoverishment/reduction in the underlying organo-mineral horizons are observed. Recognizing the humus system at the right spatial and temporal scale is crucial for the biological management of a forest. This article includes a data review, new data from a doctoral thesis, and recent comparisons of Italian and French investigations. [ABSTRACT FROM AUTHOR]

It was time to take stock. We modified the humipedon classification key published in 2018 to make it easier and more practical. This morpho-functional taxonomy of the topsoil (humipedon) was only available in English; we also translated it into French and Italian. A standardized morpho-functional classification of humipedons (roughly the top 30–40 cm of soil: organic and organomineral surface horizons) would allow for a better understanding of the functioning of the soil ecosystem. This paper provides the founding principles of the classification of humipedon into humus systems and forms. With the recognition of a few diagnostic horizons, all humus systems can be determined. The humus forms that make up these humus systems are revealed by measuring the thicknesses of the diagnostic horizons. In the final part of the article, several figures represent the screenshots of a mobile phone or tablet application that allows for a fast recall of the diagnostic elements of the classification in the field. The article attempts to promote a standardized classification of humipedons for a global and shared management of soil at planet level. [ABSTRACT FROM AUTHOR]

Soils and crops are particularly vulnerable to climate change and environmental stresses. In many agrosystems, soil biodiversity and ecosystem services provided by soils are under threat from a range of natural and human drivers. Agricultural soils are often subject to agronomic practices that disrupt soil trophic networks and make soils less productive in the long term. In this scenario, sustainable soil use aimed at improving plant/root status, growth and development plays a crucial role for enhancing the biological capacity of agricultural soils. This commentary paper is divided into the following four main sections: (i) the contentious nature of soil organic matter; (ii) soil biological quality/fertility; (iii) soil classification; and, (iv) which agricultural practices can be defined as sustainable? The published literature was analyzed within a holistic framework, with agrosystems considered as living systems where soil, vegetation, fauna and microorganisms co‐evolve and are reciprocally influenced. Ultimately, this article will suggest a better stewardship of agricultural soils as a natural capital. [ABSTRACT FROM AUTHOR]

Terms and concepts have been defined in Humusica 1, article 1 and the functioning of humus systems has been discussed in Humusica 1, article 2. Here a short overview of the matter, showing humus systems in their environment, is provided for beginners, before making field investigations. The present work is intended as a part of the field manual (HUMUSICA 1 and 2), an illustrated, easy-to-use application tool for humus systems classification, helpful even for not (yet) expert pedologists. The present article gives also a fast look at the classification, sharing Terrestrial, Histic, Aqueous and Para humus systems, every group being defined by its characteristics set in synthetic tables, and suggests a step-by-step approach allowing everyone to classify and investigate humus systems and forms. [ABSTRACT FROM AUTHOR]

In this article, we present the Terrestrial intergrades from never or only few days submersed humipedons, without permanent hydromorphic signs, until humipedons with presence of signs of partial asphyxia. Specific terms are defined and diagnostic horizons illustrated, both with the help of text explanations and photographs. The article ends with a table working as a classification field key. [ABSTRACT FROM AUTHOR]

The Special Issue Humusica 1 corresponds to a field guide for the classification of terrestrial humus systems and forms. The present first article of the issue defines vocabulary, objects and concepts necessary for: (a) field investigation, (b) understanding the process of classification, (c) assigning ecological significance to the defined morpho-functional units, (d) discussing and exchanging scientific data about humus systems. The article starts with general considerations, as the necessity humans have to classify natural objects for sharing ideas and information on them. Then the article focuses on soil as functional element of every ecosystem. Historical and recently published international definitions of soil are reported and compared to the more biological definition of the authors of the paper. Once the concept “soil” clarified, the soil profile is parted in three new sub-units, for specialised further investigations. The superficial and richer in organic matter sub-unit is labelled Humipedon. In the rest of the article, authors explain the different constituents of a general humipedon, introducing even novice soil scientists to field practice and topsoil observation. A general overview of the variety of humipedons that one may expect to find all over planet Earth, ranged in humus systems and humus forms, concludes the article. [ABSTRACT FROM AUTHOR]

This article is an as simple as possible key of classification of terrestrial (aerobic, not submersed) topsoils (organic and organic-mineral series of soil horizons). Based on the introduction exposed in Humusica 1, article 1, and using vocabulary and definitions listed in article 4, a classification is proposed for better understanding the biological functioning of the soil, partially disclosing the process of litter digestion. Five types of terrestrial topsoils, called terrestrial humus systems, are described and illustrated with the help of photographs. Within each humus system, 3–4 humus forms are also revealed, corresponding to similar series of soil horizons generated in a relatively homogeneous environment whose range of ecological factors is not so large to overstep and cause the genesis of another different humus system. The article ends with a figure that shows the relationship between Tangel and Amphi humus systems, and a dichotomous key of classification that one can easily print and bring in the field for practicing humus classification. [ABSTRACT FROM AUTHOR]

The present paper should be read after the six preceding articles dealing with fundamental concepts, vocabulary, principles and keys of our morpho-functional classification of humus systems and forms, the whole book being conceived as a guide for field studies. It now concerns seven questions which may arise when passing from knowledge to practice or, in other terms, from concepts to real things. In the field, humipedons can differ from reported photographs. Trees, bushes and/or herbs interfere with soil functioning and may generate different humipedons even over a relatively small surface. More generally a researcher must select a few unit plots representative of a given natural floor for defining the frame of his investigation. In this article, authors present some practical and theoretical landmarks and illustrate some solutions for studying humipedons in common natural sites, and replacing them within ecosystem dynamics. [ABSTRACT FROM AUTHOR]

Knowledge of a little number of specific terms is necessary to investigate and describe humipedons. This “new vocabulary” allows individuating and circumscribing particular diagnostic horizons, which are the fundamental bricks of the humipedon. Few “components” defined by specific terms characterize a specific “humipedon horizon”; few “humipedon horizons” compose a given “humus form” and some similar “humus forms” are grouped in a functional “humus system”. In this article, specific terms and humus horizons are listed and explained one by one. Field difficulties are illustrated and resolved. The aim of the article is to present in a manner as simple as possible how to distinguish in the field the soil structures allowing a morpho-functional classification of terrestrial (aerated, not submerged) humipedons. [ABSTRACT FROM AUTHOR]

Litter biodegradation is a process of life. Organisms feed, reproduce, die and decompose. Decomposition is essential, and it is never complete. In addition, the elements generated by this process become new bricks for building more complex structures in a dynamically evolving environment. In this article, we show some pictures of the main actors in litter biodegradation. We also try to associate living organisms to the soil aggregates they generate, furnishing photographs of organisms and aggregates visible in the field even with a naked eye. The transformation of dead bodies, organs or cells and droppings in the soil ecosystem is influenced by biotic and abiotic factors and hence it must be considered as a dynamic, never ending, local evolution. Instead of focusing on specific data, we have tried to present the involved phenomena to a non-specialised public (naturalists, students, teachers, etc.) through the use of graphical schemes, indicating arrows, photographs, and drawings. In the end, readers will be aware that things are not as simple as expected, that static models cannot give a precise image of a reality in constant evolution. The article can be inspected as a photo album, read as a comic strip or used as a dictionary. The authors aim to illustrate rather than to explain the relationships between humus systems, climate and biodiversity. [ABSTRACT FROM AUTHOR]

Born in Trento (Italy, 2003) for the purpose of standardising vocabulary and units of humus form classification, after publishing a first synthetic classification e-book (Zanella et al. 2011) they do not cover all site conditions in the European area. Although having basic concepts and general lines, the European (and North American, Canadian, the Humus group decided to use its classification for handling global change (Zanella and Ascher-Jenull 2018). The process is detailed in many scientific articles published in three Special Issues (Humusica 1, 2 and 3) of the journal Applied Soil Ecology. Conceptually, the whole of Humusica answers three crucial questions: A) What is soil? Soil is a biological ecosystem. It recycles dead structures and implements mineral material, furnishing more or less re-elaborated organic, mineral and organic-mineral elements to support living organisms. Article chapters: 1. Essential vocabulary; 2. Soil covers all the Earth's surfaces (soil as the seat of processes of organic matter storage and recycling); 3. Soil may be involved in the process of natural evolution (through organisms' process of recycling biomass after death). B) If soil has a biogenic essence, how should it be classified to serve such managerial purposes as landscape exploitation or protection? A useful classification of soil should consider and propose useful references to biologically discriminate soil features. Article chapters: 4. Soil corresponds to a biogenic structure; 5. TerrHum, an App for classifying forest humipedons worldwide (a first attempt to use a smartphone as a field manual for humus form classification). C) How can this soil classification be used for handling the current global change? Using the collected knowledge about the biodiversity and functioning of natural (or semi-natural) soil for reconstructing the lost biodiversity/functioning of heavily exploited or degraded soils. Article chapters: 6. Agricultural soils correspond to simplified natural soils (comparison between natural and agricultural soils); 7. Organic waste and agricultural soils; 8. Is traditional agriculture economically sustainable? Comparing past traditional farm practices (in 1947) and contemporary intensive farm practices in the Venice province of Italy. [ABSTRACT FROM AUTHOR]

The article is structured in six sections. A first portion is dedicated to the state of the art concerning climatic change and agriculture. Internet available IPCC maps and cartographic documents made by scientific Centres of research were used for illustrating forecasted climatic changes. In Sections 2 and 3 , bibliographic evidences were collected for supporting a vegetation and soil co-evolution theory. Humus, soil and vegetation systems are presented at planet level in many synthetic maps. In Sections 4 , 5 and 6 the authors discussed the human influence on the soil evolution during the Anthropocene. It appears that humans detected and used the Mull humus systems all over planet Earth for crop production and pasture. Human pressure impoverished these humus systems, which tend to evolve toward Amphi or Moder systems, losing their natural biostructure and carbon content. [ABSTRACT FROM AUTHOR]

Agro humus systems correspond to agricultural humipedons. Specific terms and diagnostic horizons are defined and topsoil profiles are described. With this new vocabulary, which focuses on the biological structure of the A horizon, we can compare these humipedons with their original natural expression. Under human pressure for agricultural purposes, natural humipedons evolve toward anthropic Mull (See Humusica 1, articles 5 and 6 for natural humus horizons) or Amphi (Mull with layers of still not incorporated OH organic horizons) humus systems. In each humus system, we define a few humus forms in order to estimate the biological health of the soil, with a rapid observation realisable in the field by the naked eye. Histic anthropic Anmoor is also possible, as well as Hydro or Epihistic intergrades to Terrestrial systems. WRB qualifiers and Soil Taxonomy subgroup adjectives are reported at the end of the article and can be used for a more detailed classification. [ABSTRACT FROM AUTHOR]

Techno humus systems correspond to man-made topsoils under prominent man influence. They may be purposely conceived for supporting agricultural activities or dumping of waste products, sometimes abandoned to an unknown evolution. Both categories needed a more scientific frame. This is the reason we classified them as morpho-functional humus systems. Improving agricultural soils with organic waste products is an ancestral practice. We present four examples of techno humus systems purposely created for supporting plant growth. Considering a simple home-made composting pile, we give a few basic notions about the biological functioning of these artificial humus systems. Humipedon functioning and structuration are similar to those observed in natural humus systems. Using even animal manure, we illustrate how to manage larger compost piles of waste for application in farming areas. Composting waste that contains animal proteins needs a more careful measurement of the temperature of the pile and a longer period of elevated temperature in the core of the pile. Mulching of pruning residues is presented in a large urban context. The use of mulch must take into account quality and composition of woody material. The lack of nutrients in some residues has to be compensated by a moderate use of appropriate mineral fertilizers. Municipal solid waste, anaerobic digestion residues (grape remains) and spent mushroom compost, eventually mixed with mineral fertilizers, have been tested in horticulture. Benefits and drawbacks are listed for each experiment, with the evolution of carbon storage along 8 years of horticultural practice. Finally, we present an example of “dump” humus system. Mine tailing wastes represent a huge problem in many countries. Pointing on their microbial activity, we show that they must be seen as manageable living humipedons, not as piles of inert rocky material. [ABSTRACT FROM AUTHOR]

Abstract: The impact of geology, climate, soil type and vegetation on forest humus forms was studied in the Veneto Region (northern Italy). A total of 352 study sites were compared by Redundancy Analysis (RDA). Humus forms were described by the structure (micro-, meso-, or macro-aggregated) of the organo-mineral A horizon, by the thickness of litter horizons and by their nomenclature, which followed the morpho-functional classification recently proposed for inclusion in the WRB-FAO. The size of aggregates within the A horizon was distributed along a common gradient embracing geology, climate, soils and vegetation. Macro-aggregation (as opposed to micro-aggregation, meso-aggregation being intermediate) was favored by carbonated (as opposed to silicated) parent rocks, warmer climate associated to lower elevation, lower soil acidity, deciduous (as opposed to coniferous) forest vegetation and relatively high clay content. The amphi group of humus forms, associated with carbonated substrates in Esalpic and Mesalpic climate districts, was distributed according to thickness of litter horizons along a gradient of soil stoniness. Biological reasons for the observed environmental influences on the size of soil aggregates, a criterion of humus form classification, were discussed to the light of knowledge on annelid (earthworm and enchytraeid) ecology. Humus forms can be easily identified and classified on the field, using a table included in the article. Our results can be used for mapping the distribution of forest humus forms in the Veneto Region, implying a better understanding of carbon cycling processes in the frame of present-day global warming. [Copyright &y& Elsevier]

Abstract: The present review was undertaken to add more information on the place taken by humus forms in plant–soil interactions. Three questions were asked: (i) are humus forms under the control of plant–soil relationships, (ii) are humus forms the main seat of these relationships, and (iii) can humus forms explain interactions between aboveground and belowground biodiversity. Some conflicting views about humped-back models of species richness may be resolved by considering a limited number of stable humus forms (here considered as ecosystem strategies) which should be treated separately rather than in a single model. Mull, moder and mor pathways are each characterized by a fine tuning between aboveground and belowground communities, the humus form (including litter) being the place where resonance between these communities takes place, both in functional and evolutionary sense. [Copyright &y& Elsevier]

Summary: The common development of vegetation and soil is a central issue in plant succession. We hypothesized that areas of woody vegetation decay and accumulation on the ground (zones of destruction or ‘micro-chablis’) played a role in the successional development of vegetation patches on tropical inselbergs and that disturbance events could be inferred from the analysis of the organic matter accumulated along a successional gradient. The study was conducted in French Guiana (South America). Nine humus profiles (each comprised of a varying number of layers) were selected in shrub thickets (∼1acre each) representative of three vegetation types of the rock savanna: canopies of pure Clusia minor (Clusiaceae), C. minor in mixture with Myrcia saxatilis (Myrtaceae) and zones of destruction. Using a dissecting microscope, a count point optical method for small soil volumes was employed to measure the volume ratio of each kind of humus component (107 categories) in the 62 layers sampled. Micro-morphological data were analysed by correspondence analysis (CA). Humus profiles varied with canopy tree type and revealed traits of past as well as trends for future plant succession. The lack of OL and OF horizons, and the presence of charred material differentiated the zones of destruction from other humus profiles and confirms the impact of spatially limited fires or lightning strikes in the cyclic development of vegetation patches. [Copyright &y& Elsevier]

Abstract: The endogeic earthworm Pontoscolex corethrurus (Glossoscolecidae) is a peregrine species commonly found in tropical lands cleared by man for cultivation. We compared the charcoal consumption and casting activity of a population of P. corethrurus from a cultivated area under repeated slash-and-burn (fallow population) with that of a population from a field cultivated after recent burning of a mature forest (forest population). Their cast production was measured in containers in the presence of pure charcoal, soil of fallow and forest origin, or a mixture of charcoal and soil. The forest population defecated less in pure charcoal than in forest soil, whereas the reverse was observed for the fallow population. When living in fallow soil, both populations defecated more at the surface of a mixture of charcoal and soil than at the surface of pure soil (×2 and ×3 with fallow and forest populations, respectively). In forest soil, both populations showed an increased charcoal consumption (×12). In the light of these experiments, we hypothesized that an adaptation of P. corethrurus to charcoal and fallow soil exists, supporting the observed distribution of this earthworm species in tropical open lands. [Copyright &y& Elsevier]

Humus forms are the seat of most biological transformations taking place in terrestrial ecosystems, being at the interface between plants, animals and microbes. The diversity of terrestrial humus forms (mor, moder and mull) can be attributed to the existence of different patterns (strategies) for the capture and use of resources by ecosystems, in ascending order of biodiversity and bioavailability. Arguments are found in the parallel development of humus forms and terrestrial ecosystems, in exclusion mechanisms between organisms living in different humus forms, and in palaeontological studies. The diversification of terrestrial life forms in the course of Earth history, concomitant with an improvement in resource availability due to the development of sedimentary layers at the surface of continents, may explain the successive appearance of more active humus forms enabling the co-existence of an increasing number of organisms. Contradictory reports about the relationships between biodiversity and stability of ecosystems can be explained by the existence of different belowground pathways making ecosystems more stable. [Copyright &y& Elsevier]

Recommends humus index as an integrated tool for the assessment of forest floor and topsoil properties. Correlation between the index and various morphological and chemical variables describing forest floors and topsoil profiles; Potential application in bulk assessment of organic matter accumulation.

Laboratory experiments were conducted using intact collembolan communities, exposed to Madit D®, a phenylurea herbicide (active ingredient isoproturon). Effects were investigated using two distinct humus types, an acid Dysmoder and a neutral Eumull. Within two weeks, no effect of the herbicide was displayed by the Eumull population, while the Dysmoder population was stimulated. When animals were able to escape from the herbicide through a perforated wall separating two compartments filled with natural soil, the behavior of collembolan communities exhibited interactive (non-additive) effects of humus type and herbicide application. The combination of an acid soil (supposedly providing greater tolerance to organic pollutants) with a neutral soil, increased biodiversity of Collembola, but caused the disappearance of some acido-sensitive species, pointing to complex relationships between pesticides, soils and soil organisms. Parallel experiments with single species demonstrated that at the recommended dose Madit D® may cause avoidance effects, but no toxicity. [Copyright &y& Elsevier]

Humusica 1 and 2 Applied Soil Ecology Special issues are field guides for humipedon classification. Contrary to other similar manuals dedicated to soil, the objects that one can describe with these guides are living, dynamic, functional, and relatively independent soil units. This is the reason to why the authors dedicated the whole article number 2 to functional considerations even before readers could go in the field and face the matter to be classified. Experienced lectors can overstep many of the sections reported in this article. If the titles of sections “1 A functional classification", "2 What is a humus system?"and "3 Energetic considerations in terrestrial systems” stimulate the reader’s curiosity, then we suggest to pass through them. Otherwise, only section “4 Climatic, plant litter, or nutritional constraints?” is crucial. Readers will understand how the soil works in terms of litter and Carbon accumulation, which one(s) among climatic, vegetational, or geological factors that intervene and strongly affect the formation processes of terrestrial (oxygenated) soils. The article concludes with a debate about a tergiversated question: can temperature influence humus decomposition? Preceding statements were used for explaining how the biological soil net can store in the soil a maximum of energy in the form of SOM, by raising a plateau partially independent of climatic conditions. [ABSTRACT FROM AUTHOR]

Abstract: The morpho-functional classification of humus forms proposed in a previous issue by Zanella and collaborators for Europe has been extended and modified, without any change in diagnostic horizons, in order to embrace a wide array of humus forms at worldwide level and to complete and make more effective the World Reference Base for Soil Resources. For that purpose 31 Humus Form Reference Groups (HFRGs) and a set of prefix and suffix qualifiers are proposed, following the rules erected for the WRB. An exhaustive classification key, respecting the principles of WRB, is suggested and examples of classification are given for some already well known humus forms. [Copyright &y& Elsevier]

Abstract: The aim of our study was to describe succession related changes in humus profiles on moderate slopes of a tropical inselberg (Nouragues, French Guiana). Nine humus profiles were collected in a stratified manner under two main communities on well-drained sites: carpets of Pitcairnia geyskesii (Bromeliaceae) and shrub thickets of Clusia minor(Clusiaceae), the latter including two stages of its dynamic development. The 53 sampled layers were analysed by an optical method, the volume ratio of 109 classes of litter/humus components being quantified by a count point method. Correspondence analysis (CA) revealed marked differences among humus forms. Pitcairnia carpets were characterized by the dominance of cyanobacteria which formed crusts with low faunal activity, except when they were colonized by enchytraeid worms. With advancing succession, we observed that leaf litter did not accumulate but rather was incorporated into organo-mineral excrements of macro-invertebrates under C. minor. The late developmental stage of Clusia thickets, characterised by the establishment of Myrcia saxatilis (Myrtaceae), showed a thick layer of undecayed litter and near absence of organo-mineral aggregates. The humus form varied from mor in cyano-bacterial crusts to tropical moder (with a few mull features) in Clusia thickets, but comparisons among humus profiles revealed more complex successional processes than expected on the basis of the composition of plant and soil animal communities. [Copyright &y& Elsevier]

Philosophy can overlap pedology. It is not casual that life begins and finishes in the soil. We separated the concepts of Humipedon, Copedon and Lithopedon. Some sections were dedicated to the founders of the movement for a new type of agriculture (agroecology). They simply proclaim to accompany the process of natural evolution instead of spending a lot of energy in hunting competitor organisms with pesticides or boosting the soil with mineral fertilisations and tillage. The core of the article is built on a biological concept of soil and shows researches supporting this view. After pointing to the soil structure and illustrating its natural genesis, explaining which cultural conditions may improve its quality, we finished the article with economic considerations, combining at planet level a program of soil restoration with a greenhouse effect mitigation. What a reader should have in mind at the end of the article: soil organisms have a prominent positive influence on soil structure and fertility; their mass is proportional to the soil organic matter quantity; it is possible to contrast the climate warming using the soil as sink of C. We estimated that the Agro Humipedons of a European economically active region could sink about 13 or 20% of its emissions, switching from conventional to minimum or no tillage during the coming 40 years. At planetary level, a well programmed 4 per 1000 action can even be more efficacious and compensate a part of the global greenhouse gas effect. [ABSTRACT FROM AUTHOR]

The aim of this article is to present a sort of state of the art from a soil classification point of view. How far are soils from their original natural state? A few years ago, the two most important soil classification systems added a classification of anthropogenic soils. Even so, the concept of anthropogenic soil differs slightly in the two systems, it is easy to recognize these soils all around us, in both urban and agricultural contexts. After presenting the main types of anthropogenic soil, using the WRB or Soil Taxonomy characteristics, we will focus on anthropogenic humipedons, presenting a table where they are subdivided into two new units called Agro (natural topsoils used for agricultural purposes) and Techno (human-made humus systems). This allows to conceptually compare soil and humipedon classifications, and prepares the field for further investigations on Agro (Humusica 2, art. 15) and Techno (Humusica 2, art. 16) humipedons. [ABSTRACT FROM AUTHOR]

Abstract: We monitored the carbon isotope composition of bulk leaves and specific long-chain alkanes during a four-year litterbag experiment using 13C-labelled leaves and unlabelled reference leaves of the European beech tree (Fagus sylvatica L.). Whereas the isotope composition of alkanes from 13C-enriched leaves exhibited a marked decrease in 13C-content, the isotope composition of unlabelled reference leaves remained nearly constant. We interpreted this difference as evidence for a microbial contribution to the long-chain alkane pool of the decomposing leaves and related it to the progressive invasion of leaves by soil organisms which was revealed upon microscopic examination. These results suggest that long-chain alkanes may not provide an unaltered record of organic carbon isotope composition in soils and sediments. [Copyright &y& Elsevier]

Abstract: The development of temperate deciduous and conifers forests stands usually results in accumulation of forest floor organic matter and a shift from mull to moder humus forms. It has been suggested that an increase in nutrient uptake by trees during their rapid growth phase leads to topsoil acidification, decrease in earthworm density and thereby a decrease in litter turnover. The focus of this paper was to examine if the mull-moder shift with forest ageing results from higher leaf litter production and/or lower litter decay rates. The objectives of this research were to determine (1) changes in macro-morphological properties of humus forms, leaf litter production, litter decay rates, soil nutrients content and pH along a 130-year pure beech (Fagus sylvatica L.) chronosequence in Normandy (Northwest France), (2) if humus form varied from mull to moder with increasing stand age, and (3) if a shift from mull to moder resulted from increased litter production, decreased litter decay rates, or both. Annual litter production did not change significantly along the chronosequence (mean 2.41tha−1). In contrast, litter decay rates decreased significantly during the rapid growth phase of trees. In consequence, the litter turnover time (1/k) was lower in the youngest stands (20 months) compared to the oldest ones (31 months). Even in the absence of a significant pattern of variation, litter production was positively correlated with the thickness of the OF (Oi) horizon. In contrast, litter decay was strongly negatively correlated with maximum thickness of the OH (Oa) horizon, suggesting that the appearance of the humification layer was mainly due to a decrease in litter decay rate. We did not find significant changes in the main properties of the organo-mineral horizon, suggesting that soil nutrient availability may not directly affect litter dynamics. We concluded that moder development along the chronosequence resulted in decreasing litter decay rates during the aggradation phase while litter production was stable. Further studies are required to identify the ecological factors responsible for moder development along forest ageing. [Copyright &y& Elsevier]

Abstract: We sampled moss and vascular forest vegetation in five ancient beech forests from northwest France, embracing in each a wide array of environmental conditions. Indirect (PCA) and direct (RDA) gradient analysis were used to discern local and regional ecological factors which explain the observed variation in species composition. Our results point to a global factor encompassing a large array of soil and light conditions, unravelled when local particularities of studied forests are singled out. The humus form, numerically expressed by the Humus Index, explains a large part of the observed variation in ground vegetation. Our study confirmed opposite trends in vascular and moss species richness according to humus condition. Ecological factors to which vascular and moss forest species respond at the regional level can be estimated directly in the field by visually inspecting humus forms and vegetation strata despite of the confounding influence of local factors. [Copyright &y& Elsevier]