Biology And Evolution Of Ferns And Lycophytes Pdf

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About once every decade for the last 80 years, a volume has been produced that reports comprehensively on recent developments in pteridology. Some have an emphasis on particular aspects, such as phylogeny, cytogenetics or gametophytes, while others have a broader coverage.

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Biology and Evolution of Ferns and Lycophytes

A survey of ferns and lycophytes of the Coastal Region of the state of Rio Grande do Sul CRRS was performed based on field work and collections of the main regional herbaria. The following were evaluated for each species: preferential habits terrestrial, epiphytic or aquatic , geographic distribution patterns and habitats forest, grassland, and wetland. The forest habitat harbored the greatest number of species , while grasslands had the fewest Our results demonstrate a strong north-to-south reduction in species richness in the study area, which is related to environmental conditions along the latitudinal gradient and, especially, microclimatic differences in the transition zone between the Atlantic Forest and Pampa biomes.

In the meridional portion of South America the "Southern Cone" , centers of species diversity and endemism, as well as limits to floristic distributions, have been related to paleogeographical alterations that produced climatic barriers and interrupted floristic continuity Sota ; Ponce et al.

At a local scale, climatic factors, such as temperature and humidity Kessler ; Kessler et al. The presence of a center of diversity and endemism of ferns and lycophytes in the Atlantic Forest of southeastern Brazil Tryon , is relevant for the diversity of southern Brazil, including the Coastal Region of the state of Rio Grande do Sul.

This mosaic is determined mainly by edaphic conditions, but also by wind action and extreme insolation Rambo However, there is general transition in vegetation cover from it being scarce and sparse at the coastline, to gradually giving rise to forests with tropical characteristics in the continental interior Rambo ; ; Klein ; Waechter Through this inventory, we seek to understand the distribution patterns of ferns and lycophytes among different vegetation physiognomies and along a latitudinal gradient in the study area, and the relationships of these plants with adjacent plant communities.

The CRRS is limited to the west, continental, side by the geomorphological formations of the "Sulriograndense" Plateau a. The origin of the Coastal Region is discussed by Tomazelli et al. Despite the relative uniformity of these coastal areas, microtopographic variation allows for two main soil types: well drained sandy soils at low elevations, and poorly drained sandy, clayish or peaty soils in the lowlands.

Most of the terrain is covered by Solodic Planosol derived from alluvial and lacustrine depositions or sandstone decomposition. These Planosol areas are poorly drained and have low fertility, with forest growing on peat.

The northern portion is characterized by Hydromorphic Humic Quartz-Soil type Neosol of sedimentary origin formed during the Holocene and featuring a high accumulated-carbon content and poor drainage. Also in the northern portion, shallow and highly fertile Chernosols are present, with rock mixtures in the A horizon.

Red Latosol can also be found, constituting deep soils, rich in iron and nutrient poor in the A horizon. Dune banks of depositional origin prevail along the coastline, composed of quartz sands of low pedogenetic structuration, low fertility and high wind mobility Moser ; Embrapa These climate conditions occur in most regions with altitudes below m alt. Field work was conducted between March and December in 19 municipalities Fig. Spot inventories were conducted in different vegetation formations with random sampling following the procedure described by Filgueiras et al.

The samples were collected and preserved following Windisch All exsiccates from the study area were annotated with updated identifications and nomenclature, and their location and habitat information were added to our database.

The taxonomic delimitation of families and genera follows Smith et al. For the genus Blechnum L. For Tryonia J. Prado, A. Cochran we followed Cochran et al. Moran et al. Species habits were categorized as terrestrial, epiphytic or aquatic and assigned to habitats: forest, grassland, wetland or combinations thereof. Holarctic: wide distribution in the temperate regions of the Northern Hemisphere and Eurasian region;. Neotropical-Antarctic: distribution in the Neotropical region, with eventual occurrence in temperate South-American regions;.

The latitudinal distributions of the taxa within the study area were verified by their presence or absence in five latitudinal ranges, according to Waechter This analysis was processed by the Past 3. A total of 3, herbarium records were evaluated, including collected during our field work and 2, previously existing in collections. In total the records represented species in 79 genera and 28 families, of which 17 were species of lycophytes and were species of ferns Tab.

The largest values for species richness were for the families: Pteridaceae 34 , Polypodiaceae 29 , Thelypteridaceae 23 , Dryopteridaceae 22 , Aspleniaceae 18 and Blechnaceae The genera with the largest values for species richness were: Thelypteris 22 , Asplenium 17 , Blechnum 12 , Elaphoglossum nine , Pecluma eight , Pleopeltis seven and Doryopteris seven , with the remaining genera varying from one to six species Tab. Richness differed among the evaluated latitudinal ranges, with the number of taxa gradually decreasing from north-to-south with Range I having species, while Range V had only Of the total species, only 21 occurred throughout the entire study area Tab.

Vegetation: F - forest; G - grassland; W - wet land. Some families such as Lygodiaceae, Marattiaceae and Tectariaceae were restricted to the extreme north. Others, such as Lindsaeaceae, Hymenophyllaceae, Lomariopsidaceae, Selaginellaceae, Dicksoniaceae and Schizaeaceae, occurred exclusively in the three northern-most latitudinal ranges Fig. With an increase in latitude, there was an increase in terrestrial species richness and a reduction in epiphytes.

Note the reduction in the number of species from north-to-south in the CRRS. Some species were found both in forest and grassland habitats 15 spp. Nine species were present in wetland environments, and an equal number of species occurred in all three environments forest, grassland and wetland.

Blechnum brasiliense, Thelypteris conspersa and Thelypteris metteniana, occurred in forest and wetland habitats, whereas Blechnum schomburgkii and Palhinhaea cernua were found in grassland and wetland environments Tab. Differences in preference patterns for vegetation formations were identified over the studied profile Fig. The species Deparia petersenii and Nephrolepis cordifolia are amphipacific; Osmunda regalis and Osmundastrum cinnamomeum are Holarctic; and Blechnum cordatum, Lophosoria quadripinnata, Hymenophyllum caudiculatum and Equisetum giganteum are Neotropical-Antarctic.

Only 21 species were common to all areas Tab. In the Neotropics, plant species richness is more strongly correlated with precipitation than edaphic variation Gentry In the Atlantic Forest, patterns of floristic change of arboreal vegetation along latitudinal gradients have been attributed to variation in temperature and rainfall distribution.

The same authors verified that rainfall and temperature increments influence processes of tropicalization and changes in vegetation types, mostly when associated with increased latitude.

For Atlantic Forest ferns and lycophytes, variables such as precipitation and evapotranspiration determine plant species composition in distinct vegetation types, as also discussed by Gasper et al.

Large-scale changes in diversity related to latitudinal gradients have been previously reported for ferns and lycophytes by Moran , with several examples of the concentration of species richness in tropical regions with accentuated decreases towards both poles.

According to this author, in South America, particularly in the tropical region, fern and lycophyte richness can reach 2, species, while in temperate regions it is only 79 species.

Sato points out that in Japan, species richness decreases with increasing latitude, mostly due to climatic conditions. Through the southern portion of CRRS, there was a reduction of the floristic richness of most families and genera. In addition to the reduction of fern and lycophyte total diversity, the incidence of species with Neotropical-Atlantic and Neotropical-Paranaense geographical distribution patterns was strongly affected. Taxa that maintain their distributions along the profile 21 spp.

The extended distribution of many tropical species in the CRRS can be interrupted by alterations to the north-to-south forest continuity. The transition from tropical rain forest to other vegetation types such as grasslands, peat forests, sandy forests, seasonal forests and riparian habitats Waechter can affect distribution patterns and inhibit dispersal of ferns and lycophytes.

These formations develop on the extreme southern slopes of the "Serra do Mar" and on the eastern face of "Serra Geral". Due to the climatic conditions of temperature and precipitation, as well as the environmental heterogeneity of vegetation and substrates, the development of larger forests with tropical species are allowed, as is the dispersal of northern floristic components Rambo ; Klein ; Waechter ; Waechter These characteristics of environment heterogeneity are fundamental to increasing fern and lycophyte diversity in the Atlantic Forest Windisch ; Dittrich et al.

Sota considers that the Pampa, along with the Chaco, are barriers separating the center of fern and lycophyte diversity in southern Brazil from the Argentinian and Chilean centers of diversity. This same reduction in diversity has been reported previously by other authors. Rambo ; ; ; observed structural and floristic modifications to vegetation along the Coastal Region, attributing the decrease in species richness to the dispersal processes that occurred throughout the region he referred to as "Porta de Torres" "Torres migrational gate".

This latitudinal gradient was later observed by Waechter unpublished data in a study of vascular epiphytes, who found a Of the 61 genera he studied, only four are found in Uruguay and one in the province of Buenos Aires Argentina.

Among the factors that influence floristic change in the north-to-south direction of the CRRS, Waechter highlights climatic discontinuity along the region. The author noted that the decrease in rainfall and temperature going south causes an increase in the number of cold days and annual frost events.

This, in turn, results in changes to vegetation communities, causing insularization and discontinuity of the arboreal component, thus interfering in the dispersal routes of tropical plant species. Ranges IV and V were observed. These results can be associated with the high dispersal capacity of fern and lycophyte spores, thereby finding niches with favorable conditions for their establishment and reproduction Tryon ; Wolf et al.

Another north-to-south floristic change in the CRRS can be seen in the reduction of epiphytic species in Range V with the concomitant prevalence of terrestrial species Fig. Beyond the need for adequate ecological conditions Sota ; Benzing , there is the requirement of host plant availability for the establishment of epipytic species Waechter unpublished data.

Some species restricted to Range I, such as Vandenboschia radicans, Lomariopsis marginata, Mickelia scandens, Polybotrya cylindrica, Olfersia cervina, are hemiepiphytes, that begin development on the ground and later climb host plants. Decreased amounts of rainfall and the existence of a drier season in the extreme south can limit the establishment and development of fern and lycophyte flora.

Ranal a points out that fern spore germination, as well as the development of the gametophyte and sporophyte, are strongly affected by desiccation. Watkins et al. Other features inherent to the reproductive success of many ferns and lycophytes can be decisive in their establishment in particular habitats.

Although this group of plants is capable of dispersing spores over long distances Holttum and can form spore banks in the ground Ranal , the spores may not encounter the microclimatic temperature, humidity and luminosity and biotic conditions mycorrhizal associations needed for germination Williams ; Sheffields Furthermore, gametophytes, even when possessing adaptations to desiccation and variable conditions of pH, soil nutrition, temperature and luminosity Farrar et al.

Despite the fact that some fern species have strategic adaptations for independent gametophytic reproduction apogamy, clonal reproduction by gemmae, pH metamorphosis the presence of water is still a requirement for the successful sexual reproduction of the gametophyte of most fern and lycophyte species Page According to Holttum , the majority of fern and lycophyte spores germinate in forest environments, and to occupy other environments with adverse conditions they need specific adaptations.

The occurrence of ferns in wetland and grassland environments is also related to expansion of these vegetation formations in the north-to-south direction, with the expansion of water bodies in wetland-region lagoons and natural grasslands most frequent in Range V.

For example, Blechnum brasiliense has adapted to a variety of environments that differ in their edaphic composition, vegetation formations and microclimate conditions Silva et al. Data compiled by Nimer and Moreno , point out clear differences in the climatic patterns of the two extremes of CRRS.

Furthermore, the average number of days with frost increases from three to 15 days in the extreme south Range V. This southern, more temperate climate can explain the presence of species with Neotropical-Antarctic or Holarctic distribution patterns in the floristic composition of Range V, such as Osmunda regalis and Equisetum giganteum. According to these authors, frost affects fern sporophytes in three ways: direct mortality by freezing caused by wind, soil freezing, and hydric unavailability caused by ice.

In order for this species to occur in these regions of low temperatures and frost, adaptations at all development stages are required. In northeastern United States, gametophytes of the genus Vittaria Sm.

Sehnem suggested that the low richness of ferns and lycophytes in some regions of the state of Rio Grande do Sul is influenced by the reduced amount of time for the migration of additional species recent paleogeographic evolution and, consequently, the limited distribution of these species in the extreme south.

This phenomenon may explain the decrease of fern and lycophyte species in the temperate floras of the "Southern Cone", which can have three to four times fewer species than tropical floras such as in southeastern Brazil Ponce et al. The impoverishment of the extreme southern flora of CRRS may also be related to past paleoenvironmental conditions.

Virtual issue: Ecology and evolution of pteridophytes in the era of molecular genetics

Sutherland, B. Barrett, J. Beck, M. Latvis, M. McKain, E. Sigel , and N.


Cambridge Core - Evolutionary Biology - Biology and Evolution of Ferns and Lycophytes. Frontmatter. pp i-vi. Access. PDF; Export citation 7 - Evolution of the nuclear genome of ferns and lycophytes. pp By Takuya Nakazato.


The system can't perform the operation now. Try again later. Citations per year. Duplicate citations. The following articles are merged in Scholar.

With their team of contemporary scholars, the editors present a thorough coverage of fundamental topics necessary for obtaining an uptodate understanding of the biology of ferns and lycophytes.

Skip to search form Skip to main content You are currently offline. Some features of the site may not work correctly. DOI: Ranker and C. Ranker , C. Haufler Published Biology.

The past quarter-century has witnessed a revolution in our understanding of the phylogenetics, systematics, and ecology of pteridophytes ferns and lycophytes , particularly due to the rapid accumulation of plastid sequence data and a renewed interest in the ecology of the sexual phase of the life cycle. We here compile 19 papers recently published in the Journal of Plant Research dealing with the biology of pteridophytes, grouped into six categories: 1 breeding systems, 2 species complexes and polyploidization, 3 fossil taxa, 4 gametophyte ecology, 5 systematics, 6 biodiversity. We hope this collection of papers will be of value to researchers interested in this fascinating group of plants. Our understanding of the phylogeny of pteridophytes ferns and lycophytes was revolutionized starting in with the sequencing and phylogenetic analysis of the plastid rbcL gene Hasebe et al. This ushered in the modern era of pteridophyte systematics, which has witnessed increasingly sophisticated analyses of the evolutionary relationships of seed-free vascular plants Pryer et al.

It will provide essential background information for those beginning research in any of the areas covered. This book fills a major gap in the literature of the past decade and provides a modern overview of the biology and evolution of these fascinating plants. Tom A. Ranker received his Ph. Christopher H.

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1 Comments

  1. Cookie-Singh 15.01.2021 at 17:35

    A survey of ferns and lycophytes of the Coastal Region of the state of Rio Grande do Sul CRRS was performed based on field work and collections of the main regional herbaria.