Klaus Mehltreter

Klaus Mehltreter
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Klaus verified their affiliation via an institutional email.
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Klaus verified their affiliation via an institutional email.
  • Dr. rer. nat.
  • Senior Researcher and Professor at Institute of Ecology INECOL

About

128
Publications
95,032
Reads
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2,596
Citations
Introduction
I am an ecologist currently working at the Institute of Ecology (INECOL) in Xalapa, Mexico. My research is focused on the ecology of ferns (including lycophytes). With my students and collaborators, we investigate the biodiversity of Mexican ferns, their dispersal by spores, their germination in the wild and in vitro, their adaptations to climate (phenology), susceptibility to disturbance and role as indicators. A current project is focussed on their interactions with herbivorous insects and mites.
Current institution
Institute of Ecology INECOL
Current position
  • Senior Researcher and Professor
Additional affiliations
September 1998 - present
Institute of Ecology INECOL
Position
  • Professor
Description
  • Lab courses: Bioinformatics I and II, Scientific writing; Field courses: Fern ecology, Comparative plant ecology
March 1996 - September 1998
Comisión de Investigaciones Científicas de la Provincia de Buenos Aires
Position
  • Researcher
May 1990 - December 1996
Ulm University
Position
  • Research Assistant
Editor roles

Publications

Publications (128)
Article
Full-text available
Ferns have been considered ecological indicators of soil nutrient composition and can be adapted to limestone (calcicole), volcanic substrates (calcifuge), or both (generalists). However, how many species exhibit substrate preferences and how these substrates affect their leaf nutrient composition remains unclear. We studied the occurrence of fern...
Article
The few current metabolomic studies on ferns are mostly restricted to a single species or focused on specific compounds. We performed an untargeted metabolomic study on six of the most common fern species from the cloud forest, followed by a targeted analysis of 64 phenolic compounds, many of which have been associated with herbivore defense. The u...
Article
Limestone substrates cover 20–30% of the continents and are the habitat of many calcareous fern species, which grow preferably or exclusively on this substrate. In this review, we resume the limestone properties and their implications for calcareous ferns. Additionally, we investigate the taxonomic distribution of limestone ferns by comparing five...
Article
Premise: Extrafloral nectaries are mainly studied in angiosperms, but have also been reported in 39 fern species. Here we provide a global review of nectaries in ferns, and study their structure, function, and nectar sugar composition in two genera. Methods: We searched in the literature and living plant collections of botanical gardens for indi...
Article
Full-text available
Background and Aims: The mutualism between leaf-cutting ants and the fungus Leucoagaricus gongylophorus is a remarkable instance of insect-fungus symbiosis. In this study, we aimed to make a molecular identification of the L. gongylophorus strain obtained from the fungal garden of Atta mexicana, to compare the mycelial growth of the strain in sever...
Article
Full-text available
Soil and water characteristics in micro basins with different land uses/land cover (LULC) can influence riparian vegetation diversity, stream water quality, and benthic diatom diversity. We analyzed 18 streams in the upper part of the La Antigua River basin, México, surrounded by cloud forests, livestock pastures, and coffee plantations. Concentrat...
Article
Full-text available
Urbanization is creating a new global biome, in which cities and suburbs around the world often resemble each other more than the local natural areas they replaced. But while urbanization can profoundly affect ecology at local scales, we know little about whether it disrupts large-scale ecological patterns. Here we test whether urbanization disrupt...
Article
Recent investigations have revealed the coordination and trade-offs between above- and belowground traits in structuring functional strategies of plant communities, but none of these have addressed ferns, a diverse plant lineage in tropical forests. We investigated terrestrial ferns from the perspective of how below- and aboveground traits are coor...
Article
Full-text available
The relative importance of allochthonous and autochthonous carbon (C) as sources of energy for tropical stream food webs remains an open question. Allochthonous C might be the main energy source for small and shaded forest streams, while autochthonous C is more likely to fuel food webs draining land uses with less dense vegetation. We studied food...
Preprint
Full-text available
Urbanization can profoundly disrupt local ecology. But while urban areas now stretch across latitudes, little is known about urbanization’s effects on macroecological patterns. We used standardized experiments to test whether urbanization disrupts latitudinal gradients in seed predation, a macroecological pattern that shapes community assembly and...
Article
Full-text available
Questions Leaf traits play an important role in the epiphyll colonization of leaves. Although ferns are common in forest understories, they have been widely ignored as possible host plants. We asked: How does epiphyll colonization (cover, richness, and composition) differ among fern species depending on their leaf longevity and surface wettability?...
Article
In several montane forests around the world, epiphytes coexist in mats, sharing the rhizosphere and forming histosol‐type soils rich in nutrients. The role of these epiphytes in the formation of canopy soil and the fitness costs that epiphytes face when cohabiting in these mats are unknown. In a lower montane cloud forest in central Veracruz, Mexic...
Article
Full-text available
During fern spore germination, lipid hydrolysis primarily provides the energy to activate their metabolism. In this research, fatty acids (linoleic, oleic, palmitic and stearic) were quantified in the spores exposed or not to priming (hydration–dehydration treatments). Five fern species were investigated, two from xerophilous shrubland and three fr...
Article
Full-text available
The Cretaceous–Cenozoic expansion of tropical forests created canopy space that was subsequently occupied by diverse epiphytic communities including Eupolypod ferns. Eupolypods proliferated in this more stressful niche, where lower competition enabled the adaptive radiation of thousands of species. Here, we examine whether xylem traits helped shape...
Article
Cloud forests are highly diverse mountain ecosystems, but habitat loss and fragmentation threaten many species with extinction. Demographic parameters are essential to implement effective conservation actions and to understand the mechanisms of species coexistence, but there are very few population models with transition matrices for fern species....
Article
Ferns have been exposed to herbivorous insects since the latter evolved in the Devonian. Currently, ferns suffer similar percentages of leaf herbivory as angiosperms. Therefore, they often use a combination of chemical defenses as protection against herbivores. In this review, we summarize the distribution of five groups of biomolecules that may ac...
Article
Background and Aims Ferns are the second largest group of vascular plants and are distributed nearly worldwide. Although ferns have been integrated into some comparative ecological studies focussing on hydathodes, there is a considerable gap in our understanding of the functional anatomy of these secretory tissues that are found on the vein endings...
Article
Premise: Terminal velocity (Vt) is one main factor in biological particles dispersal but is scarcely studied for anemochorous fern spores. Moreover, the effect of spore characteristics on Vt has not been evaluated yet. In the present study, we measured the Vt of 1234 spores of 21 fern species and two Selaginella microspores with a video image analy...
Article
Paramos are highly diverse ecosystems of tropical mountains above the tree line, where plant communities are frequently exposed to extreme climatic conditions. In this study, we analyzed the influence of microenvironmental factors (slope, cover of rocks, herbs and shrubs) on the richness, abundance and composition of fern and orchid assemblages in...
Article
Fern‐feeding insects have not received much attention because ferns have been considered herbivore‐resistant plants. However, ferns are attacked by herbivorous insects, including specialized spore feeders. For a better understanding of host specialization, we investigated the effect of experimentally transferred larvae of a spore‐feeding moth, Stat...
Chapter
The approximately 12,000 extant fern and lycophyte species comprise about 4% of the vascular plant diversity, and still ca. 40–50 new species are described each year. Although ferns possess a nearly worldwide distribution, they are especially abundant in tropical mountains at mid-elevations and on remote oceanic islands. The Tropical Andes and Sout...
Article
Full-text available
While most plants die below a threshold of water content, desiccation-tolerant species display specific responses that allow them to survive extreme dehydration. Some of these responses are activated at critical stages during water loss and could represent the difference between desiccation tolerance (DT) and death. Here, we report the development...
Article
Full-text available
Ferns and lycophytes are the second and third largest lineages of vascular plants, yet our understanding of their interactions with phytophagous insects is very limited. In this study, we reviewed herbivorous insects, their feeding habits and host preferences on these two plant groups, searched for any evidence of coevolution, and discussed possibl...
Article
Full-text available
Some fern species are among the world’s worst weeds, but little is known about the establishment of alien ferns in relatively conserved ecosystems. Exotic species may outperform native ones during germination and facilitate invasion into new environments. Macrothelypteris torresiana is an Asian fern species that was introduced and spread throughout...
Article
• Host plant selection by herbivores is driven by a complex array of cues, including leaf traits and previous leaf damage. Herbivore-associated cues to host selection at the plant and leaf scale aid understanding of mechanisms responsible for host preference that might translate into increased performance, as well as processes structuring herbivore...
Preprint
Full-text available
Plants can tolerate some degree of dehydration but below a threshold of water content most plants die. However, some species display specific physiological, biochemical, and molecular responses that allow survival to desiccation. Some of these responses are activated at critical stages during water loss and could represent the difference between de...
Preprint
Full-text available
Ferns are the second largest lineage of vascular plants, yet our understanding of their interactions with phytophagous insects is very limited. Which insects feed on ferns? Do insects prefer specific fern taxa? Which feeding habit is the most common on ferns? Is there any evidence for coevolution between insects and ferns? Is our current knowledge...
Article
Full-text available
Opportunistic diversification has allowed ferns to radiate into epiphytic niches in angiosperm dominated landscapes. However, our understanding of how ecophysiological function allowed establishment in the canopy and the potential transitionary role of the hemi‐epiphytic life form remain unclear. Here, we surveyed 39 fern species in Costa Rican tro...
Preprint
Full-text available
Opportunistic diversification has allowed ferns to radiate into epiphytic niches in angiosperm dominated landscapes. However, our understanding of how ecophysiological function allowed establishment in the canopy and the potential transitionary role of the hemi-epiphytic life form remain unclear. Here, we surveyed 39 fern species in Costa Rican tro...
Article
In the laboratory, non-chlorophyllic (orthodox) fern spores are long-lived. However, their potential longevity might not reflect ecological longevity, which determines their viability within the natural soil spore bank. Our aim was to compare ecological (in a semi-natural soil bank) and potential (storage in laboratory–suboptimal conditions) longev...
Method
Full-text available
This script is the result of teaching a postgraduate course over the last ten years about writing and publishing scientific articles in English.
Article
Full-text available
Vascular plants exhibit defense syndromes, a variety of interdependent defensive traits against herbivores, which may considerably differ between plant groups. Although ferns are an abundant component of tropical forest understories, studies of fern–herbivore interactions are scarce, and none has focused on the underlying defense syndromes. To exam...
Article
Full-text available
Atta mexicana is a polyphagous insect that can exploit a wide range of plant species to cultivate its main food source, the symbiotic fungus Leucoagaricus gongylophorus. In the present study, we evaluated the foraging rate of A. mexicana workers among leaves of three favored and three rejected plant species under laboratory conditions. In addition,...
Article
We report two new records of rare fern species for the state of Morelos, Mexico: Hemionitis pinnatifida and Myriopteris longipila subsp. brevipila. For both species, we describe their main characteristics, cite studied material and present photographs and distribution maps, followed by comments on their habitat characteristics, historical record, r...
Article
• Canopy soil (CS) volume reflect epiphyte community maturity, but little is known about the factors that retain CS or species succession within it. Humus fern species (e.g. Phlebodium areolatum) appear capable of retaining CS. • In ten Quercus spp. we sampled 987 epiphyte mats to examine the role of the common epiphyte species and crown traits det...
Poster
Full-text available
Version 1 - Español - Spanish
Article
Full-text available
Plants of the humid tropics seal their leaf surfaces with hydrophobic waxes to keep them dry and clean by repelling water and diminishing the adhesion of spores of pathogenic and epiphyllous organisms. We investigated the chemical composition and micromorphological structure of waxes of the upper leaf surfaces of six terrestrial (Diplazium drepanol...
Article
Full-text available
Species interactions have long been predicted to increase in intensity toward the tropics and low elevations because of gradients in climate, productivity, or biodiversity. Despite their importance for understanding global ecological and evolutionary processes, plant-animal interaction gradients are particularly difficult to test systematically acr...
Article
Full-text available
Fern–insect interactions have not received the same attention as angiosperm–insect interactions have. It has even been stated that ferns may have very few interactions with animals because of their lack of flowers, fruits, and seeds. Consequently, for many decades fern–insect interactions have been overlooked and underestimated, especially for high...
Preprint
Full-text available
Fern-insect interactions have not received the same attention as angiosperm-insect interactions have. It has even been stated that ferns may have very few interactions with animals because of their lack of flowers, fruits and seeds. Consequently, for many decades fern-insect interactions have been overlooked and underestimated, especially for highl...
Article
Fern spores and seeds initiate germination with fast water uptake, followed by a stationary phase with no appreciable water uptake and biochemical and metabolic processes that precede germination. After that, seed, germination is avoided by dehydration, as part of the priming treatments. After dehydration, seeds maintain their metabolic advances (h...
Article
Full-text available
Although leaf-cutter ants of the genus Atta are polyphagous, they discriminate between food plant species. Ferns represent the second largest group of vascular plants and are especially abundant in tropical forest understories, but seem to be avoided as food plants. We studied the leaf damage caused by leaf-cutter ants and its relation with water c...
Article
Full-text available
Forest edges typically exhibit higher luminosity and lower humidity than the forest interior, resulting in an abiotic gradient. However, the degree of abiotic difference can be affected from the type of the matrix, influencing the selection of species. We compared the floristic and phytosociological structure of understory communities of ferns and...
Article
Full-text available
Forest edges typically exhibit higher luminosity and lower humidity than the forest interior, resulting in an abiotic gradient. However, the degree of abiotic difference can be affected from the type of the matrix, influencing the selection of species. We compared the floristic and phytosociological structure of understory communities of ferns and...
Article
Full-text available
The edge effect impacts the richness, composition, and abundance of plant species, as a result of environmental changes caused by forest fragmentation. Ferns are considered important indicators of environmental quality because they are directly affected by these alterations. The purpose of this study was to analyze the influence of the edge effect...
Preprint
Full-text available
Species interactions have long been predicted to increase in intensity towards the tropics and low elevations, due to gradients in climate, productivity, or biodiversity. Despite their importance for understanding global ecological and evolutionary processes, plant-animal interaction gradients are particularly difficult to test systematically acros...
Article
Full-text available
Ferns adapt their phenological characteristics such as leaf production, leaf fertility and leaf mortality to the habitat conditions optimizing their opportunities to grow, reproduce, and disperse. The leaf phenology of the herbaceous, dimorphic fern Danaea geniculata was studied in a Brazilian submontane tropical forest and compared with several ot...
Article
The new tree fern species Cyathea leoniae from Northern Peru is described, illustrated and differential characteristics from most similar species are provided.
Article
Full-text available
Herbaceous plants rely on a combination of turgor, ground tissues and geometry for mechanical support of leaves and stems. Unlike most angiosperms however, ferns employ a sub-dermal layer of fibers, known as a hypodermal sterome, for support of their leaves. The sterome is nearly ubiquitous in ferns, but nothing is known about its role in leaf biom...
Data
A pair-wise correlation matrix of the log-transformed data constructed with the “psych” package (Revelle, 2017) in the R environment. The panels present scatter plots of variable pairs in which the x axis in each plot is the column variable, and the y axis the row variable. Each scatter plot shows the loess locally fit regression with an elllipse a...
Data
Non-significant relationships between the percentage of sclerenchyma that occupies the petiole and flexural rigidity (A) and leaf area (B).
Data
A list of Pteridaceae species, Genbank atpA and rbcL sequence ID numbers, and source attributions.
Data
Additional petiolar cross-sections from the surveyed Pteridaceae taxa.
Article
The dispersal of diaspores is a key process for successful reproduction, survival and evolution of plants. To model the dispersal of biological propagules (e.g., spores, pollen) it is necessary to measure their terminal velocity (Vt) in air. In this study, we used a new method based on video image analysis to measure Vt of spores of seven fern taxa...
Article
Full-text available
Premise of the study: This paper investigates the occurrence and evolution of aluminum (Al) accumulation within ferns and lycophytes, which is characterized by Al concentrations above 1000 mg·kg(-1) in aboveground plant tissues. We hypothesize that this feature is more common in ferns than in angiosperms, and potentially correlated with growth for...
Chapter
Full-text available
The ongoing degradation, fragmentation and loss of natural forests, makes management activities for ecological restoration a priority. To recover the structure and function of these natural ecosystems, basic scientific knowledge is needed urgently. Minimal restoration intervention (passive restoration) aims to halt anthropogenic disturbance and to...
Article
The knowledge of spore mass and size is fundamental to our understanding of dispersal of airborne spores and the biogeography of fern species. Thus, in the present study we present a simple method to measure mass of biological particles; the spore mass of 23 fern species was measured and related with spore size and type. Species could be grouped ac...
Article
Premise of research. Structure-function relationships of xylem in tropical fern species remain poorly explored despite the taxonomic diversity and ecological importance of ferns in tropical environments. The aim of this study is to broaden our understanding of how tropical fern morphology and anatomy are influenced by light availability. Methodolog...
Article
Glyphosate, a systemic herbicide, is used against weeds in agricultural fields as well as against invasive plants in pastures, forest plantations, and urban environments. Its frequent and widespread use can negatively impact the surrounding natural non-target vegetation following the accidental drift of spray droplets, leaching, or persistence as r...
Article
Full-text available
http://www.biodiversidad.gob.mx/Biodiversitas/Articulos/biodive119art2.pdf
Article
Neotropical bracken fern invades disturbed forests and burned and abandoned pastures in Latin America, inhibiting the growth of associated vegetation and altering community structure. Cutting of all aboveground vegetation every 6 to 12 mo has proven to be inefficient as a control method. We studied the impact of selective cutting of bracken every 2...
Conference Paper
Full-text available
For the last 250 years, fern research was primarily focused on systematics, taxonomy and morphology and more recently on molecular phylogenetics as well as floristic and ethnobotanic studies. Only about 4% of annual fern publications are indexed as ecological studies (Mehltreter & Hooper 2013). Fern ecology is however experiencing an increasing int...
Book
Full-text available
This book includes an explanation about the culture of epiphytes that grow in shadow coffee plantations in central Veracruz, previously occupied by cloud forest.
Article
Full-text available
Invasive species that alter ecosystems are often successful competitors due to their effects on nutrient cycling. Sphaeropteris cooperi (Cyatheaceae; Australian tree fern) has been invading intact Hawaiian rainforests for decades and displacing the dominant native tree fern Cibotium glaucum (Cibotiaceae). S. cooperi produces more leaves that grow f...
Article
Full-text available
leaf lifespan (llS) is a fundamental ecological trait of special importance to growth and survival of ferns. the llS of ferns varies between two and 49 months depending on species and site conditions. Whereas temperate ferns are mainly summergreen, tropical ferns are mostly evergreen. however, some tropical ferns have seasonal or even deciduous lea...
Article
Full-text available
Agriculture, Ecosystems and Environment j o u r n a l h o m e p a g e : w w w . e l s e v i e r . c o m / l o c a t e / a g e e Benefits and costs of epiphyte management in shade coffee plantations a b s t r a c t While epiphytes contribute to the biodiversity and structural complexity of shade coffee, their removal from the shade trees is a common...
Chapter
The contribution gives a commented list to the most relevant bibliography published on Fern and Lycophyte Ecology. It is available to individual or institutional subscribers online.
Article
Full-text available
In ferns, leaf traces in the petioles are sometimes surrounded by a circumendodermal band (CB), a cell layer of varying structure and arrangement. We studied the CB of 89 fern species from 53 genera and 28 families, and its correlation with leaf trace configuration, leaf dissection and life form, to explore its anatomical diversity, possible origin...
Article
Full-text available
Ferns have been screened for decades in search of antibacterial, antiviral or antifungal components useful for medicine. However, bioassays with living test organisms to study the ecological significance of the secondary chemistry of plants have been scarce. We experimented with leafcutter ants, which are generalist plant consumers to cultivate a s...
Article
Full-text available
En Veracruz, con base en la Flora de México de Mickel y Smith (2004), nuevas especies recién descritas y una revisión puntual en los herbarios MEXU, NY, UAMIZ y XAL para confirmar la correcta determinación y presencia de ciertas especies mencionadas en trabajos taxonómicos anteriores, existen 562 taxa (557 especies, tres variedades y dos híbridos;...
Chapter
Full-text available
1. Extant ferns and lycophytes are ecologically important and contribute 4% of the vascular plant diversity on Earth but currently face an unprecedented threat caused mainly by human disturbances such as fire or land use change. Few fern species benefit from these disturbances, and most become less abundant or locally extinct. 2. Current risk asse...
Article
The ecology of ferns is a rapidly growing discipline that offers new and exciting insights into general ecological principles and applications. Progress has been made in studying fern biogeography, population dynamics, natural resource use, disturbance responses, species interactions and links with humans (Table 10.1). In this concluding chapter, w...
Article
Ferns immediately capture the imagination of all who are fortunate enough to notice them. With their large, highly dissected and shiny green leaves, ferns are so visually appealing that many are sold as ornamentals. Most moist woodlands will have a number of fern species blanketing the understory with their pungent foliage. In tropical woodlands, f...
Article
Biogeographical patterns of ferns and angiosperms are the result of a combination of vicariance and long distance dispersal, but due to their more effective dispersal via spores, the latter is more frequent among ferns. Therefore, fern species tend to have wider ranges and the relative number of fern species compared with seed plants is highest on...
Article
1. Ferns and lycophytes have developed a wide spectrum of antagonistic and mutualistic relationships with fungi and animals. While some of these interactions, such as endomycorrhizae, are old and may have coexisted with their host plants for a long time, other interactions may have originated more recently, such as some herbivorous insects that hav...
Article
Full-text available
We studied the structure of newly found entomogenous galls in three Indian species of Selaginella, and the biochemical changes during gall development as well as their seasonal abundances in 10 squares of 1.5 m under two differing habitat conditions, open and covered. Adult wasps (Cynipidae) initiate two types of gall formation by oviposition, sphe...
Book
Ferns are an integral part of the world's flora, appreciated for their beauty as ornamentals, problematic as invaders and endangered by human interference. They often dominate forest understories, but also colonize open areas, invade waterways, and survive nutrient-poor wastelands and eroded pastures. This is the first comprehensive summary of fern...
Article
Se describe e ilustra Psychotria perotensis Castillo-Campos perteneciente al subgénero Heteropsychotria Steyermark (Rubiaceae, Psychotrieae) como una especie nueva de la región La Cortadura, Coatepec, en las faldas orientales del volcán Cofre de Perote, Veracruz, México. Esta especie muestra similitudes con P. phanerandra (Standley & Steyermark) Lo...
Article
Tropical montane cloud forests deliver important goods and services to society, such as timber, the supply and purification of fresh water, and carbon sequestration. In spite of their relevance, current deforestation rates are very high, at the expense of affecting the provision of ecosystem services. We explore the impact of land use change in ter...
Article
Full-text available
Ponthieva brenesii (Orchidaceae) and Piper xanthostachyum (Piperaceae) are reported for the first time in the State of Veracruz, Mexico. The first species is very rare, and has a restricted distribution range previously limited within Mexico to Cuernavaca, Morelos and Teziutlán, Puebla. The second species is a climbing herb, uncommon growth habit i...
Article
Full-text available
Ponthieva brenesii (Orchidaceae) and Piper xanthostachyum (Piperaceae) are reported for the first time in the State of Veracruz, Mexico. The first species is very rare, and has a restricted distribution range previously limited within Mexico to Cuernavaca, Morelos and Teziutlán, Puebla. The second species is a climbing herb, uncommon growth habit i...
Article
Full-text available
En la región centro de Veracruz existen fragmentos de bosque mesófilo de montaña (BMM) que no se han explorado. Nosotros estudiamos la composición fl orística y algunos aspectos de la estructura de un BMM en la localidad de La Cortadura. El muestreo de leñosas, herbáceas y epífitas vasculares se realizó en tres microcuencas (1000 m2 cada una). Las...
Article
Full-text available
In the central area of Veracruz a few remnants of cloud forest (CF) have not been explored previously. We studied the floristic composition and some structural features of one of these remnants of CF in La Cortadura. Sampling of woody plants, herbs and vascular epiphytic species was made in three river basins (plots of 1000 m2 each). The sampled ar...
Article
Full-text available
In the central area of Veracruz a few remnants of cloud forest (CF) have not been explored previously. We Studied file floristic composition and some structural features of one of these remnants of CF in La Cortadura. Sampling of woody plants, herbs and vascular epiphytic species was made in three river basins (plots of 1000 m(2) each). The sampled...

Questions

Questions (2)
Question
I am aware about image-analysing software listed at http://www.plant-image-analysis.org/,
and do not refer neither to public image databases such as Pl@antNet or iNaturalist to compare pictures or ask others for help nor to applications providing interactive keys such as LucID,
but automized software which analyses pictures of leaves and suggests possible identifications.
Question
Leaf litter movement may depend not only on the inclination of the slope, but air and rainfall, soil humidity, leaf size, plant species, surface cover (bare soil, mosses). My interest is focused on the question how long bare soil is covered by leaf litter and consequently how much time propagules may have to wait for their chance, if they require direct light for germination.  

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