No abstract
Florisitic ground surveys in tropical rain forests are laborious and time consuming, so we tested to what degree reflectance differences visible in Landsat Thematic Mapper (TM) satellite images can be used to predict differences in florisitic composition and species richness among rain forest sites. To gain ecological understanding of the rain forest ecosystem, we also tested to what extent variation in these vegetation characteristics can be explained by edaphic site conditions. The study was conducted in a relatively homogeneous area of Amazonian rain forest in Yasuní National Park, Ecuador. We established 27 transects of 5 m × 500 m within an area of ∼20 km × 25 km to study edaphic and floristic patterns mainly within the tierra firme (non‐inundated) forest. In each transect, soil samples were collected for chemical and textural analyses, and the abundance of each species belonging to two understory plant groups, pteridophytes (ferns and fern allies) and the Melastomataceae, was assessed. Floristic similarity between transect pairs varied widely and ranged from almost no overlap in species composition to very high overlap. The among‐transect floristic similarity patterns of the two plant groups were strongly correlated with each other no matter whether presence–absence or abundance data were used. The floristic similarity patterns were also strongly correlated with the similarity in pixel values of the infrared bands in the Landsat TM satellite image and with the similarity in most of the measured soil variables. Similarity in species richness, on the contrary, was neither correlated with similarity in pixel values nor with similarity in most of the soil variables. We conclude that reflectance patterns in satellite images can be efficiently used to predict landscape‐scale floristic and edaphic patterns in tierra firme rain forest. Predicting patterns in species richness, on the other hand, is not possible in the same straightforward manner. These results have important practical implications for land use and conservation planning as well as for ecological and biodiversity research. Corresponding Editor: C. A. Wessman.
We documented the floristic composition of pteridophytes (ferns and fern allies) and Melastomataceae in Yasuní National Park, Amazonian Ecuador. Our main questions were: (1) Are the density of individuals, species richness, and/or species diversity (measured with Shannon's H′) of the two plant groups related to edaphic differences? and (2) How many of the pteridophyte and Melastomataceae species are non–randomly distributed in relation to a soil base content gradient within terra firme (non–inundated forest). To answer these questions, we sampled 27 line transects of 500 × 5 m distributed in an area of ca 20 × 25 km. The study area included a permanent 50 ha plot established to monitor forest dynamics; thus, our results also provide information on landscape–scale floristic variability to which results from within the plot can be compared. A total of 45,608 individuals and 140 species of pteridophytes, and 4893 individuals and 89 species of the Melastomataceae, were counted in the transects. Both with pteridophytes and with Melastomataceae, a clear negative correlation was found between the amount of extractable bases in the soil and the number of plant individuals encountered in a transect. With Melastomataceae, species richness and species diversity also were negatively correlated with soil base content, but with pteridophytes they were not. More than 50 percent of the common species of both pteridophytes and Melastomataceae were nonrandomly distributed along the soil cation content gradient within terra firme. We conclude that while the species richness patterns observed in one plant group are not indicative of similar patterns in other plant groups, it seems likely that a substantial (but unknown) proportion of species belonging to other plant groups will be found to show distribution patterns that reflect edaphic preferences within terra firme forests.
To better understand changes in the distribution and diversity within plant functional types along an elevational gradient and the potential mechanisms driving such changes, we studied species richness of ferns at six elevations along a forested elevational gradient in Costa Rica, from La Selva Biological Station at 30 m a.s.l. up the slopes of Volcán Barva to 2960 m a.s.l. Among the samples from all the sites combined, we found 264 species from 60 genera. Sixty-nine species were terrestrial, 113 were canopy epiphytes, and 121 were low-trunk epiphytes. Only one species occupied both canopy and terrestrial habitats at any of the study sites. Overlap of canopy and low-trunk species composition was relatively low (18%), and lower still was the overlap of terrestrial and low-trunk species (12%). Total species richness peaked at the 1000-m site for canopy and low-trunk epiphytic species. In contrast, the richness of terrestrial species rose to a mid-elevation maximum and remained relatively constant at higher elevations.In an effort to explain elevational patterns of species richness, we examined mean annual rainfall and temperature, light intensities in the canopy and at ground level, and the mid-domain effect. Of the explanatory factors examined, the mid-domain effect accounted for most of the elevational pattern. We found little evidence that environmental gradients drive patterns of fern species richness on this spatial scale.
Leaves are lateral determinate structures formed in a predictable sequence (phyllotaxy) on the flanks of an indeterminate shoot apical meristem. The origin and evolution of leaves in vascular plants has been widely debated. Being the main conspicuous organ of nearly all vascular plants and often easy to recognize as such, it seems surprising that leaves have had multiple origins. For decades, morphologists, anatomists, paleobotanists, and systematists have contributed data to this debate. More recently, molecular genetic studies have provided insight into leaf evolution and development mainly within angiosperms and, to a lesser extent, lycophytes. There has been recent interest in extending leaf evolutionary developmental studies to other species and lineages, particularly in lycophytes and ferns. Therefore, a review of fern leaf morphology, evolution and development is timely. Here we discuss the theories of leaf evolution in ferns, morphology, and diversity of fern leaves, and experimental results of fern leaf development. We summarize what is known about the molecular genetics of fern leaf development and what future studies might tell us about the evolution of fern leaf development.
Despite the extraordinary significance leaves have for life on Earth, their origin and development remain vigorously debated. More than a century of paleobotanical, morphological, and phylogenetic research has still not resolved fundamental questions about leaves. Developmental genetic data are sparse in ferns, and comparative studies of lycophytes and seed plants have reached opposing conclusions on the conservation of a leaf developmental program. We performed phylogenetic and expression analyses of a leaf developmental regulator (Class III HD-Zip genes; C3HDZs) spanning lycophytes and ferns. We show that a duplication and neofunctionalization of C3HDZs probably occurred in the ancestor of euphyllophytes, and that there is a common leaf developmental mechanism conserved between ferns and seed plants. We show C3HDZ expression in lycophyte and fern sporangia and show that C3HDZs have conserved expression patterns during initiation of lateral primordia (leaves or sporangia). This expression is maintained throughout sporangium development in lycophytes and ferns and indicates an ancestral role of C3HDZs in sporangium development. We hypothesize that there is a deep homology of all leaves and that a sporangium-specific developmental program was coopted independently for the development of lycophyte and euphyllophyte leaves. This provides molecular genetic support for a paradigm shift in theories of lycophyte leaf evolution.
Homosporous vascular plants are typically depicted as extreme inbreeders, with bisexual gametophytes that produce strictly homozygous sporophytes. This view is promulgated in textbook life cycles despite ample evidence that natural populations of most species regularly outcross. We review research on a variety of mechanisms, including genetic load, asynchronous production of eggs and sperm, and pheromonal control of gamete production, that actively promote heterozygosity in ferns and lycophytes. Evolution of the land plants cannot be reconstructed without accurate depictions of the unique life cycle that has helped make ferns the second most diverse lineage of vascular plants on Earth. With revised illustrations and definitions, we provide scientists, educators, and students with a contemporary understanding of fern and lycophyte reproduction, revealing them as evolutionarily dynamic and exploiting a wide range of mating systems.
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