ABSTRACT
PREMISE: Hybridization is recognized as an important mechanism in fern speciation, with many allopolyploids known among congeners, as well as evidence of ancient genome duplications. Several contemporary instances of deep (intergeneric) hybridization have been noted, invariably resulting in sterile progeny. We chose the christelloid lineage of the family Thelypteridaceae, recognized for its high frequency of both intra- and intergeneric hybrids, to investigate recent hybrid speciation between deeply diverged lineages. We also seek to understand the ecological and evolutionary outcomes of resulting lineages across the landscape. METHODS: By phasing captured reads within a phylogenomic data set of GoFlag 408 nuclear loci using HybPhaser, we investigated candidate hybrids to identify parental lineages. We estimated divergence ages by inferring a dated phylogeny using fossil calibrations with treePL. We investigated ecological niche conservatism between one confirmed intergeneric allotetraploid and its diploid progenitors using the centroid, overlap, unfilling, and expansion (COUE) framework. RESULTS: We provide evidence for at least six instances of intergeneric hybrid speciation within the christelloid clade and estimate up to 45 million years of divergence between progenitors. The niche quantification analysis showed moderate niche overlap between an allopolyploid species and its progenitors, with significant divergence from the niche of one progenitor and conservatism to the other. CONCLUSIONS: The examples provided here highlight the overlooked role that allopolyploidization following intergeneric hybridization may play in fern diversification and range and niche expansions. Applying this approach to other fern taxa may reveal a similar pattern of deep hybridization resulting in highly successful novel lineages.
Subject(s)
Ferns , Genetic Speciation , Hybridization, Genetic , Phylogeny , Ferns/genetics , Ferns/classification , PolyploidyABSTRACT
The latest research shows that ferns and lycophytes have distinct evolutionary lineages. The codon usage patterns of lycophytes and ferns have not yet been documented. To investigate the gene expression profiles across various plant lineages with respect to codon usage, analyze the disparities and determinants of gene evolution in primitive plant species, and identify appropriate exogenous gene expression platforms, the whole-genome sequences of four distinct species were retrieved from the NCBI database. The findings indicated that Ceratopteris richardii, Adiantum capillus-veneris, and Selaginella moellendorffii exhibited an elevated A/U content in their codon base composition and a tendency to end with A/U. Additionally, S. capillus-veneris had more C/G in its codons and a tendency to end with C/G. The ENC values derived from both ENC-plot and ENC-ratio analyses deviated significantly from the standard curves, suggesting that the codon usage preferences of these four species were primarily influenced by genetic mutations and natural selection, with natural selection exerting a more prominent influence. This finding was further supported by PR2-Plot, neutrality plot analysis, and COA. A combination of RSCU and ENC values was used as a reference criterion to rank the codons and further identify the optimal codons. The study identified 24 high-frequency codons in C. richardii, A. capillus-veneris, and Diphasiastrum complanatum, with no shared optimal codons among the four species. Arabidopsis thaliana and Ginkgo biloba exhibited similar codon preferences to the three species, except for S. moellendorffii. This research offers a theoretical framework at the genomic codon level for investigating the phylogenetic relationships between lycophytes and ferns, shedding light on gene codon optimization and its implications for genetic engineering in breeding.
Subject(s)
Codon Usage , Evolution, Molecular , Ferns , Genome, Plant , Ferns/genetics , Ferns/classification , Selection, Genetic , Codon/genetics , Phylogeny , Base Composition/genetics , Viridiplantae/geneticsABSTRACT
A new species of Asplenium L. (Aspleniaceae) is described and illustrated. Asplenium sylvaticum is endemic of the Yungas and Paraná biogeographic provinces in the Southern Cone. The main diagnostic characters for this taxon are: apical pinnae with one to four caudate lobes at the base, similar in size to lateral pinnae, smooth pinnate blades with 18-36 pairs of caudate pinnae and spores with reticulate perispore, winged folds with scarce equinulaes on the margins and the laesurae wide with smooth margin. The new species belongs to the A. serra species complex group, characterised by xeromorphic habit, creeping rhizome and coriaceous fronds with branched scales on both surfaces, and resembles the other species inhabiting in Argentina, A. achalense and A. serra, in overall leaf architecture and shape but differs of the aforementioned species (and the other species of the A. serra complex) by the unique combination of characters of its rhizomatic scales, fronds and spores. A key with the morphological and palynological characters that differentiate A. sylvaticum from the other taxa belonging to the A. serra complex in the South American Cone and Bolivia is given.
Subject(s)
Ferns , Forests , Ferns/classification , Ferns/anatomy & histology , BrazilABSTRACT
Present-day geographic and phylogenetic patterns often reflect the geological and climatic history of the planet. Neontological distribution data are often sufficient to unravel a lineage's biogeographic history, yet ancestral range inferences can be at odds with fossil evidence. Here, I use the fossilized birth-death process and the dispersal-extinction cladogenesis model to jointly infer the dated phylogeny and range evolution of the tree fern order Cyatheales. I use data for 101 fossil and 442 extant tree ferns to reconstruct the biogeographic history of the group over the last 220 million years. Fossil-aware reconstructions evince a prolonged occupancy of Laurasia over the Triassic-Cretaceous by Cyathealean tree ferns, which is evident in the fossil record but hidden from analyses relying on neontological data alone. Nonetheless, fossil-aware reconstructions are affected by uncertainty in fossils' phylogenetic placement, taphonomic biases, and specimen sampling and are sensitive to interpretation of paleodistributions and how these are scored. The present results highlight the need and challenges of incorporating fossils into joint inferences of phylogeny and biogeography to improve the reliability of ancestral geographic range estimation.
Subject(s)
Ferns , Fossils , Phylogeny , Phylogeography , Ferns/genetics , Ferns/classification , Biological EvolutionABSTRACT
BACKGROUND AND AIMS: The staghorn fern genus Platycerium is one of the most commonly grown ornamental ferns, and it evolved to occupy a typical pantropical intercontinental disjunction. However, species-level relationships in the genus have not been well resolved, and the spatiotemporal evolutionary history of the genus also needs to be explored. METHODS: Plastomes of all the 18 Platycerium species were newly sequenced. Using plastome data, we reconstructed the phylogenetic relationships among Polypodiaceae members with a focus on Platycerium species, and further conducted molecular dating and biogeographical analyses of the genus. KEY RESULTS: The present analyses yielded a robustly supported phylogenetic hypothesis of Platycerium. Molecular dating results showed that Platycerium split from its sister genus Hovenkampia ~35.2 million years ago (Ma) near the Eocene-Oligocene boundary and began to diverge ~26.3 Ma during the late Oligocene, while multiple speciation events within Platycerium occurred during the middle to late Miocene. Biogeographical analysis suggested that Platycerium originated in tropical Africa and then dispersed eastward to southeast Asia-Australasia and westward to neotropical areas. CONCLUSIONS: Our analyses using a plastid phylogenomic approach improved our understanding of the species-level relationships within Platycerium. The global climate changes of both the Late Oligocene Warming and the cooling following the mid-Miocene Climate Optimum may have promoted the speciation of Platycerium, and transoceanic long-distance dispersal is the most plausible explanation for the pantropical distribution of the genus today. Our study investigating the biogeographical history of Platycerium provides a case study not only for the formation of the pantropical intercontinental disjunction of this fern genus but also the 'out of Africa' origin of plant lineages.
Subject(s)
Phylogeny , Phylogeography , Plastids , Polypodiaceae , Polypodiaceae/genetics , Polypodiaceae/classification , Plastids/genetics , Biological Evolution , Africa , Ferns/genetics , Ferns/classification , Evolution, MolecularABSTRACT
Abstract Introduction: Tree ferns are significant components of temperate, tropical and subtropical forests, contributing to shape complex forest stand structures. Objectives: 1) to describe the population structure of Cyathea costaricensis in a remnant cloud forest of West-central Mexico; 2) to characterize and relate the floristic composition and the structure of the most important tree species associated to the C. costaricensis population and; 3) to describe the environment where C. costaricensis occurs. Methods: We estimated the Importance Value Index (IVI) to select the most important canopy-dominant species associated to C. costaricensis; we constructed height and Diameter at Breast Height (DBH) frequency distributions for those selected species according to IVI as well as for C. costaricensis population; we computed the asymmetry of the frequency distributions through the coefficient of skewness and the probability density function via the Kernel density estimation. We tested for differences between canopy-dominant tree species and C. costaricensis population structure by the non-parametric Wilcoxon rank sum test. Results: C. costaricensis individuals presented the smallest heights and intermediate DBH sizes as compared with the canopy-dominant species, with statistically significant differences for height but not for DBH according to the Wilcoxon test. Most of the tree fern individuals were located in uneven terrains and over the base slope of the terrain; canopy openness and Total Radiation Under the Canopy values were similar to those reported for Cyathea species elsewhere. Conclusions: We confirm the hypothesis of comparable structure between the canopy-dominant species and the C. costaricensis population only for DBH; on the contrary, for trunk height, there were statistically significant differences; the small heights of C. costaricensis suggest their coexistence in the understory through sheltering from the taller canopy-dominants. Mostly all individuals of C. costaricensis were confined to local environmental conditions, particularly to physiography.
Resumen Introducción: Los helechos arborescentes son componentes importantes de los bosques templados, tropicales y subtropicales, que contribuyen a dar forma a rodales de estructuras complejas. Objetivos: 1) Describir la estructura poblacional de Cyathea costaricensis en un remanente de bosque nuboso del centro-oeste de México; 2) Caracterizar y relacionar la composición florística y la estructura de las especies arbóreas más importantes asociadas a la población de C. costaricensis y; 3) Describir el ambiente donde se encuentra C. costaricensis. Métodos: Estimamos el Índice de Valor de Importancia (IVI) para seleccionar las especies dominantes más importantes del dosel asociadas a C. costaricensis; para las especies seleccionadas según IVI, construimos distribuciones de frecuencia de alturas y diámetros a la altura del pecho (DAP), así como para la población de C. costaricensis. Calculamos la asimetría de las distribuciones de frecuencia a través del coeficiente de asimetría y la función de densidad de probabilidad mediante la estimación de densidad de Kernel. Probamos las diferencias entre las especies de árboles dominantes en el dosel y la estructura de la población de C. costaricensis mediante la prueba no paramétrica de suma de rangos de Wilcoxon. Resultados: los individuos de C. costaricensis presentaron las menores alturas y tamaños intermedios de DAP en comparación con las especies dominantes del dosel, con diferencias estadísticamente significativas para la altura, pero no para el DAP según la prueba de Wilcoxon. La mayoría de los individuos de helechos arborescentes se ubicaron en terrenos irregulares y sobre la pendiente baja del terreno. Los valores de apertura del dosel y Radiación total bajo el dosel fueron similares a los reportados para las especies de Cyathea en otros lugares. Conclusiones: Confirmamos la hipótesis de que hay una estructura similar entre las especies dominantes del dosel y la población de C. costaricensis solo para el DAP; por el contrario, para la altura del fuste, hubo diferencias estadísticamente significativas; las pequeñas alturas de C. costaricensis sugieren su coexistencia en el sotobosque a través de la cobertura árboles dominantes del dosel. La mayoría de los individuos de C. costaricensis fueron encontrados confinados a condiciones ambientales locales, en particular a la fisiografía.
Subject(s)
Ferns/classification , MexicoABSTRACT
Abstract We analyzed floristic variations in fern's metacommunity at the local scale and their relationship with abiotic factors in an Atlantic Forest remnant of northeastern Brazil. Floristic and environmental variations were accessed on ten plots of 10 × 20 m. We performed cluster analyses, based on Bray-Curtis dissimilarity index to establish the floristic relationship. The influence of abiotic factors: luminosity, temperature, relative air humidity and relative soil moisture was evaluated from a redundancy analysis. We found 24 species belonging to 20 genera and 12 families. The fern's flora showed high floristic heterogeneity (>75% for most of the plot's associations). The fern's metacommunity was structured along an abiotic gradient modulated by temperature, luminosity, and relative soil moisture.
Resumo Analisamos as variações florísticas na metacomunidade de samambaias em escala local e sua relação com fatores abióticos em um remanescente de Floresta Atlântica no Nordeste do Brasil. Variações florísticas e ambientais foram acessadas a partir de dez parcelas de 10 × 20 m. Realizamos análises de cluster, baseado no índice de similaridade de Bray-Curtis para estabelecer relações florísticas. A influência de fatores abióticos: luminosidade, temperatura, umidade relativa do ar e umidade relativa do solo foram avaliadas a partir da análise de redundância. Encontramos 24 espécies pertencentes a 20 gêneros e 12 famílias. A flora de samambaias exibiu uma elevada heterogeneidade florística (>75% para a maioria das associações entre plots). Observou-se que a metacomunidade de samambaias estava estruturada ao longo de um gradiente abiótico modulado pela temperatura, luminosidade e umidade relativa do solo.
Subject(s)
Ferns/classification , Biodiversity , Environment , Brazil , Adaptation, Physiological/physiology , Cluster Analysis , ForestsABSTRACT
RESUMO A espécie Dicksonia sellowiana, conhecida popularmente como xaxim, é uma samambaia natural do continente americano e encontrada no Brasil na Mata Atlântica. Em 2001 sua utilização para confecção de vasos para jardinagem foi proibida, e a mesma foi inserida na lista do IBAMA como espécie ameaçada de extinção. O presente trabalho descreve o efeito de diferentes técnicas de extração, incluindo ultra-som, turbólise, maceração, infusão, decocto e soxhlet utilizando etanol e solvente hidroalcoólico 70%, sob aspectos físicos e químicos. Foi avaliado o rendimento de cada extração, o perfil por CLAE-DAD, o conteúdo de polifenóis, flavonoides, proantocianidinas e esteroides além das atividades antioxidantes pelos métodos DPPH e redução do complexo fosfomolibdênio. Os dados foram avaliados estatisticamente através de desvio padrão, Tuckey e PCA. Os resultados demonstraram que a extração por Soxhlet utilizando ambos os solventes é a que possui maior rendimento final, porém o rendimento não está diretamente relacionado com o aumento nos teores de bioativos totais e atividades antioxidantes. As extrações por decocção e turbólise utilizando solvente hidroalcoólico foram as que apresentaram melhores resultados nos teores totais e na atividade antioxidante, resultados que ficam evidentes quando avaliados por PCA, demonstrando que este solvente e estas extrações são mais seletivas no estudo proposto.
ABSTRACT The species Dicksonia sellowiana, popularly known as xaxim, is a fern native to the Americas and found in Brazil in the Atlantic Forest. In 2001, its use for making gardening pots was banned, and it was added to the IBAMA [Brazilian Institute of the Environment and Renewable Natural Resources] list as an endangered species. This paper describes the effect of different extraction techniques – including ultrasound, turbolysis, maceration, infusion, decoction, and soxhlet using ethanol and 70% hydroalcoholic solvent on its physical and chemical aspects. The yield of each extraction was evaluated, as well as the profile by HPLC-DAD, the polyphenol content, flavonoids, proanthocyanidins, and steroids, in addition to the antioxidant activities through the DPPH and phosphomolybdenum complex reduction methods. The data were statistically analyzed through standard deviation, Tukey, and PCA. The results showed that the soxhlet extraction using both solvents had the highest overall yield; however, the yield is not directly related to the increase in total bioactive content and antioxidant activities. Extractions by decoction and turbolysis using a hydroalcoholic solvent exhibited the best results in total content and antioxidant activity – results that are evident when evaluated through PCA, demonstrating that this solvent and these extractions are the most selective in the proposed study.
Subject(s)
Ferns/classification , Antioxidants/analysis , SolventsABSTRACT
Cyathea atrovirens occurs in a wide range of habitats in Brazil, Paraguay, Uruguay and Argentina. In the Brazilian State of Rio Grande do Sul, this commonly found species is a target of intense exploitation, because of its ornamental characteristics. The in vitro cultura is an important tool for propagation which may contribute toward the reduction of extractivism. However, exogenous contamination of spores is an obstacle for the success of aseptic long-term cultures. This study evaluated the influence of different sterilization methods combined with storage conditions on the contamination of the in vitro cultures and the gametophytic development of C. atrovirens, in order to establish an efficient propagation protocol. Spores were obtained from plants collected in Novo Hamburgo, State of Rio Grande do Sul, Brazil. In the first experiment, spores stored at 7oC were surface sterilized with 0.5, 0.8 and 2% of sodium hypochlorite (NaClO) for 15 minutes and sown in Meyers culture medium. The cultures were maintained in a growth room at 26±1ºC for a 12-h photoperiod and photon flux density of 100μmol/m²/s provided by cool white fluorescent light. Contamination was assessed at 60 days, and gametophytic development was scored at 30, 60, 120 and 130 days of in vitro culture, analyzing 300 individuals for each treatment. There was no significant difference in culture contamination among the different sodium hypochlorite concentrations tested, and all treatments allowed for the development of cordiform gametophytes at 130 days of culture. In the second experiment, spores stored at 7 and -20°C were divided into two groups. Half of the spores were surface sterilized with 2% of NaClO for 15 minutes and the other half was not sterilized. All spores were sown in Meyers medium supplemented with one of the following antibiotics: nystatin, Micostatin® and actidione. The culture conditions and the procedures used for evaluating contamination and gametophytic development were the same described for the first experiment. No contamination was observed in spores stored at -20°C and treated with NaClO and actidione. In all treatments, cordiform gametophytes presenting antheridia were observed at 120 days. The percentages of these gametophytes increased from 120 to 130 days and no significant differences were observed among treatments. Archegonia were observed on cordiform gametophytes at 130 days. The findings provide data relevant to in vitro propagation procedures of this species, which may increase the availability of plants for ornamental purposes, therefore contributing to the reduction of the exploitation of endangered tree ferns species. Rev. Biol. Trop. 62 (1): 299-308. Epub 2014 March 01.
Cyathea atrovirens (Langsd. & Fisch.) Domin (Cyatheaceae) se presenta en una amplia gama de hábitats en Brasil, Paraguay, Uruguay y Argentina. Debido a sus características ornamentales, la especie es objeto de intensa explotación. El cultivo in vitro es una herramienta importante para la propagación lo que puede contribuir a la reducción del impacto de las actividades extractivas. Sin embargo, la contaminación exógena de esporas es un obstáculo para el éxito de cultivos asépticos a largo plazo. Este estudio evaluó la influencia de diferentes métodos de esterilización en combinación con las condiciones de almacenamiento sobre la contaminación de los cultivos in vitro y el desarrollo gametofítico de C. atrovirens. En el primer experimento, las esporas almacenadas a 7°C se esterilizaron superficialmente con 0.5, 0.8 y 2% de hipoclorito de sodio (NaClO) durante 15 minutos y se sembraron en medio de cultivo de Meyer. Aunque no hubo diferencia en la contaminación de lós cultivos entre las concentraciones de hipoclorito de sodio de las diferentes pruebas, en el tratamiento con 2% NaClO se observó un mayor porcentaje de gametofitos cordiformes a los 130 días. En el segundo experimento, las esporas almacenadas a 7 y -20°C fueron divididas en dos grupos. La mitad de las esporas se esterilizaron con 2% de NaClO durante 15 minutos y la otra mitad no fue esterilizada. Todas las esporas se sembraron en medio de Meyer suplementado con uno de los siguientes antibióticos: nistatina, Micostatin® o actidiona. No se observó contaminación de las esporas almacenadas a -20°C y tratadas con NaClO y actidiona. En todos los tratamientos, se observaron gametofitos cordiformes con anteridios y arquegonios. Los resultados proporcionan datos relevantes para la propagación in vitro de C. atrovirens, que pueden aumentar la disponibilidad de las plantas para fines ornamentales, contribuyendo así a la reducción de la exploración de las especies de helechos arborescentes en peligro de extinción.
Subject(s)
Ferns/growth & development , Germ Cells, Plant/growth & development , Germination/physiology , Sterilization/methods , Culture Media , Ferns/classification , Ferns/drug effects , Germ Cells, Plant/drug effects , Germination/drug effects , Spores/growth & development , Time FactorsABSTRACT
Although the genus Blechnum has been largely studied, problems of species characterization, nomenclature, classification and taxonomy remain unsolved. For these reasons, in this study we intend to deepen in the characterization of the arborescent group of Blechnum with a review of eight species: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare and B. werckleanum.Type material and several collections from diverse herbaria were studied. For morphological studies we used both herbarium and fresh material collected by the authors in fieldwork between 2005 and 2010. Rhizomes, stipes, vascularization of costae, as well as several characters of the lamina, such as outline, division, venation, margin structure, epidermal patterns, stomata, presence and types of hypodermis, and mesophyll of pinnae were studied. Important aspects of the morphology of the sporophyte have been analyzed, and external characters have been correlated with internal diagnostic ones to use the most reliable external traits in keys and descriptions. Taxonomy has been updated, and corrections have been introduced to the species nomenclature and comments were included to summarize the current knowledge of the geographical distribution and ecology. The presence of B. tabulare in America is confirmed, and the taxonomy of the species is updated, since external and internal morphology indicated that other related, American and African species, such as B. spannagelii and B. madagascariense must be included under that name. Similar studies carried out on B. schomburgkii indicated that the name is a synonym of B. auratum subsp. auratum, while B. auratum subsp. columbiense is treated here as a well characterized species. Detailed study of numerous specimens allowed to characterize B. moritzianum, being the correct name to apply to the recently described B. yungense. A refreshed description of the species is presented together with a list of new, selected materials to illustrate current known geographical distribution. Expanded data on ecology, geographical distribution and affinities are given, and a key for species identification is presented.
La bibliografía sobre el género Blechnum L. es muy abundante tanto en lo que respecta a las especies neotropicales como paleotropicales. Sin embargo, aún faltan estudios anatómicos completos, una caracterización actualizada de su morfología externa e interna, taxonomía, citología y afinidades. En este estudio se revisaron ocho especies de Blechnum de hábito arborescente: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare y B. werckleanum. Se analizaron caracteres de rizomas, estípites y costas, división de la lámina, y venación, estructura de márgenes, modelos epidérmicos, presencia y tipos de hipodermis y mesofilo de las pinnas. Se estudió por primera vez la hipodermis, un área dermatoide adaxial presente en el mesofilo de todas las especies. Se correlacionaron caracteres externos e internos con el fin de utilizar los externos más confiables en claves y descripciones. Los estudios realizados confirmaron nuevamente la presencia de B. tabulare en América, que incluyen en su sinonimia a B. spannagelii de América tropical y B. madagascariense de África. También se concluyó que el nombre B. schomburgkii es un sinónimo de B. auratum subsp. auratum, en tanto que B. auratum subsp. columbiense se trata aquí con rango de especie. Se caracterizó a B. moritzianum, el nombre que corresponde aplicar a la recientemente descrita B. yungense. Se presentó una descripción actualizada de las especies, se resolvieron varios problemas nomenclaturales pendientes; se presentaron datos amplios sobre la ecología, distribución geográfica y afinidades entre las especies, así como una clave para determinarlas.
Subject(s)
Ferns/anatomy & histology , Ferns/classification , Classification/methods , Terminology as TopicABSTRACT
(Estudios epidérmicos en helechos y licófitas medicinales de Provincia de Córdoba, Argentina). Se realiza un análisis epidérmico de los órganos vegetativos y reproductivos de 28 helechos y licófitas medicinales, que crecen en Córdoba (Argentina). Los objetivos fueron: a) caracterizar en detalle la epidermis de todos los órganos; b) contribuir a la diferenciación de taxones estrechamente relacionados y c) proveer información adicional para un adecuado control de calidad de los helechos medicinales que se comercializan. Se describen en detalle distintos tipos de pelos, escamas, microescamas, estructuras de respiración y características de las células epidérmicas. Para cada taxón, se provee una descripción acompañada de dibujos lineales y fotomicrografías. Se describen por primera vez pelos y escamas inéditos y se mencionan tipos de estomas novedosos en varios taxones. Desde el punto de vista taxonómico, el indumento y los tipos de estomas fueron los caracteres más significativos para diferenciar taxones. El aporte al control de calidad queda resumido en los caracteres diagnósticos de la clave y en la iconografía que permiten la identificación de una especie cuando la droga vegetal se encuentra fragmentada o pulverizada.
Subject(s)
Ferns/classification , Plant Epidermis , Plant Stomata/classification , Argentina , Photomicrography , Plants, MedicinalABSTRACT
Spores morphology and synangia in neotropical fern species of Marattia (Marattiaceae). The Marattiaceae are represented by a small family of four to six genera that bear esporogenous structures of two types: sorus with free eusporangia in Angiopteris and Archangiopteris, and indurated synangium in Christensenia, Danaea and Marattia. Marattia is a pantropical genus of about eight to ten species in the paleotropic and seven to eight species in the neotropic. In order to describe the spores and sinangia morphology, this study analyzed the shape of the receptacles, and the position of the synangia, and evaluated the spores with SEM, of seven neotropical species of the genus Marattia: M. alata, M. cicutifolia, M. excavata, M. interposita, M. laevis, M. laxa y M. weinmanniifolia from several collections. The receptacles were fully developed in M. cicutifolia and M. laevis, and scarcely overelevated in the rest of the species. The synangium was ellipsoidal and had intramarginal to supramedial position in the laminae. The spores of Marattia were elliptic. Among the taxa, only monolete spores were found, with no trilete, aborted or deformed spores. The laesura was linear and reached about two of the total length of the spore. The perispore appears as a continuous thin layer deposited on the exospore according to its ornamentation in M. cicutifolia and M. laevis. It is smooth in M. alata, rugate in M. excavata and pustulate-rugate in two species: M. interposita and M. laxa. The exospore is echinate in M. cicutifolia and M. laevis and pustulate in the other species. In M. weinmannifolia spores produced by the same sinangium may have different ornamentation types. We concluded that, while the presence of ellipsoidal and superficial synangia and monolete spores aperture were generic traits, the micro and macro-ornamentation types of the perispores and exospores vary at specific level. Besides, macro-ornamentation can be bulliform (pustulate), a combination of bulliform and muriform types (pustulate-rugate), muriform (rugate-retate) and stelliform (echinate); finally, granular micro-ornamentation can be seen frecuently in perispores. Rev. Biol. Trop. 59 (4): 1833-1844. Epub 2011 December 01.
La familia Marattiaceae, que incluye de cuatro a seis géneros, presenta estructuras esporógenas de dos tipos: sinangios en Christensenia, Danaea y Marattia; y soros con eusporangios libres en Angiopteris y Archangiopteris. Marattia es un género pantropical con unas ocho-diez especies en el paleotrópico y siete-ocho en el neotrópico. Mediante MEB se estudiaron las esporas de las siete especies neotropicales de Marattia: M. alata, M. cicutifolia, M.excavata, M. interposita, M. laevis, M. laxa y M. weinmanniifolia, basado en la forma de los receptáculos y la posición de los sinangios en material proveniente de distintas colecciones. Los receptáculos son hiperdesarrollados en M. cicutifolia y M. laevis, y apenas sobreelevados en el resto de las especies. Los sinangios son elipsoidales y la posición en la lámina es intramarginal a medial. Las esporas son elípticas, siempre monoletes y no se encuentran esporas triletes, abortadas o deformadas. La lesura tiene una longitud aproximadamente igual a la mitad del diámetro de la espora. El perisporio aparece como una capa delgada continua que se deposita siguiendo los procesos del exosporio, en M. cicutifolia y M. laevis. Es liso en M. alata, rugado en M. excavata y pustulado-rugado en dos especies: M. interposita y M. laxa. El exosporio es equinado en M. cicutifolia y M. laevis, y pustulado en las otras especies. En M. weinmanniifolia, las esporas producidas por el mismo sinangio tienen diferentes tipos de ornamentación. La presencia de sinangios elipsoidales superficiales y de esporas monoletes, son rasgos genéricos; mientras que los tipos de micro-ornamentación y macroornamentación en exosporios y perisporios son caracteres diagnósticos específicos. La macro-ornamentación puede ser buliforme (pustulada), una combinación de buliforme y muriforme (pustulada-rugada), muriforme (rugada-retiada) y esteliforme (equinada); es frecuente la presencia de micro-ornamentación granular en el perisporio.
Subject(s)
Ferns/cytology , Spores/cytology , Costa Rica , Ferns/classification , Spores/ultrastructureABSTRACT
Diversity of ferns and lycophytes from Reserva Natural Punta Lara, Buenos Aires , Argentina. Punta Lara Natural Reserve shelters the relict of the Southernmost riparian forest in the world, where the flora is threatened by the anthropic expansion and by the introduction of exotic species. In this area the ferns and lycophytes grow in particular environments as marshland, grassland and marginal forest. The aim of this work was to evaluate the diversity of ferns and lycophytes that inhabit in Punta Lara Natural Reserve. Field trips were conducted from 2006 to 2009 and local herbaria and specific bibliography were also revised. As a result of our study twenty four taxa of ferns and lycophytes were recognized. Their habit was mainly terrestrial (70%), fewer aquatic (17%) and epiphytic (13%). Five native taxa were scarcely recorded whereas two native species previously documented were not found. We assume that the main threats on ferns and lycophytes populations in Punta Lara Reserve are: removal for comercialization, invasion by exotic species and climatic fluctuations (drought periods). Keys for taxa identification are given and strategies for the conservation of threatened taxa are proposed. Rev. Biol. Trop. 59 (3): 1037-1046. Epub 2011 September 01.
Hasta el año 1987 el número de helechos y licofitas citados para la Reserva Punta Lara era de 30, incluyendo taxa específicos e infraespecíficos. El objetivo de este trabajo fue evaluar la diversidad de los helechos y licofitas que crecen en esta reserva, donde se halla el relicto más austral de las selvas en galería. Para llevar a cabo este estudio se realizaron viajes de campo durante los años 2006-2009 y se consultaron herbarios locales así como bibliografía especializada. Como resultado del estudio se determinaron 24 taxa de helechos y licofitas, de hábito predominantemente terrestre (70%); pocas acuáticas (17%) y epífitas (13%); dos especies exóticas, mientras que otras dos especies nativas citadas previamente no fueron halladas. Cinco taxa fueron registrados con escasa frecuencia. Se infiere que las principales causas de amenaza para las poblaciones de licofitas y helechos en la Reserva Natural Punta Lara son la extracción para su comercialización, la introducción de plantas exóticas y las fluctuaciones climáticas (periodos de sequía). Se presenta una clave de identificación de los taxa registrados y se plantean futuras estrategias de conservación de las especies consideradas vulnerables.
Subject(s)
Biodiversity , Ferns/classification , Argentina , Conservation of Natural ResourcesABSTRACT
We have investigated how edge effects influence the fern community of Jaguarão Forest (08º 35' 49" S and 35º 15' 39" W), located in the district of Rio Formoso, Pernambuco, Brazil. A comparative analysis was made of the interior and edge of the fragment of forest, regarding the richness, abundance and diversity of ferns in the two areas. Six plots of 10 × 20 m were chosen, three in each area. A total of 381 ferns were recorded, which were distributed among 25 species, 17 genera and 12 families. The two areas (edge and interior) were found to differ, with distinct relative air humidities and temperatures (p = 0.00254 and p = 0.00019, respectively). The interior showed higher diversity (t = 7.251 and p = 0.018) and richness (t = 6.379 and p = 0.023) than the edge area, but the same abundance (t = 1.728; p = 0.226) as the edge. Regarding the composition of the flora, it was clear that the interior is a habitat completely distinct from the edge with regard to the fern community, given that only one species, Adiantum petiolatum Desv., was common to both environments. It was concluded that the edge effect causes a decrease in richness and abundance of the fern species found in Jaguarão Forest, where the more sensitive species are being replaced by species that are tolerant to the disturbance caused by the creation of an edge.
Este trabalho verificou como os efeitos de borda influenciam as comunidades de pteridófitas presentes na Mata do Jaguarão (08º 35' 49" S e 35º 15' 39" W), município de Rio Formoso, Pernambuco. O estudo foi feito através de análise comparativa entre os ambientes de interior e de borda. Para verificar a riqueza, abundância e diversidade das pteridófitas nos dois ambientes, foram estabelecidas seis parcelas de 10 × 20 m (200 m²), sendo três para cada ambiente. Foram contabilizados 381 indivíduos de pteridófitas, distribuídos em 25 espécies, 17 gêneros e 12 famílias. Os dois ambientes (interior e borda) caracterizaram-se diferentes entre si, com umidade relativa do ar e temperatura distintas (p = 0,00254 e p = 0,00019, respectivamente). O ambiente de interior apresentou maior diversidade (t = 7,251 e p = 0,018) e riqueza (t = 6,379 e p = 0,023) que o ambiente de borda, de acordo com o teste estatístico empregado (teste t pareado). A abundância, por outro lado, foi a mesma em interiores e bordas do fragmento (t = 1,728 e p = 0,226). Quanto à composição florística, ficou claro que o interior do fragmento estudado constitui um ambiente totalmente distinto da borda, no que se refere à comunidade pteridofítica, com apenas uma espécie comum aos dois ambientes, Adiantum petiolatum Desv. Assim, o presente estudo constatou que o efeito de borda causa diminuição na riqueza e abundancia das espécies de pteridófitas ocorrentes na Mata do Jaguarão, com a existência de substituição das espécies mais sensíveis pelas espécies tolerantes aos impactos causados por esse efeito.
Subject(s)
Ecosystem , Ferns/classification , Biodiversity , Brazil , Ferns/growth & development , Population DensityABSTRACT
Morphological stomatal traits, such as size, form and frequency, have been subject of much literature, including their relationships with environmental factors. However, little effort have focused on ferns, and very few in the genus Blechnum. Stomatal length, width and frequency (as stomatal index) of a number of specimens of fourteen Neotropical species of Blechnum were measured in adult pinnae. The aim of the work was to find biometrical relationships between stomatal traits and between stomatal traits and habit, habitat and ecosystem of the plants. Statistical analyses of data were conducted using Exploratory Data Analysis and Multivariate Statistical Methods. Stomatal length and width showed a very high correlation, suggesting an endogenous, genetic control, thus giving these traits a considerable diagnostic utility. With respect to the relationships between stomatal traits and environment, we found significant statistical relationships between altitude and stomatal index. We also addressed the interpretation of the ecological- selective significance of various assemblages of stomatal traits in a diverse conjunction of habits, habitats and ecosystems. Rev. Biol. Trop. 59 (1): 403-415. Epub 2011 March 01.
Los caracteres morfológicos estomáticos, tales como tamaño, forma y frecuencia, han sido objeto de abundante investigación, incluyendo su relación con los factores ambientales. Sin embargo, poco esfuerzo se ha realizado en esta materia en helechos y menos todavía en el género Blechnum. En este trabajo se midieron la longitud, anchura y frecuencia (como índice estomático) de estomas de pinnas adultas de un número de individuos en catorce especies de Blechnum neotropicales. El objetivo fue encontrar relaciones biométricas entre los caracteres estomáticos, y entre los caracteres estomáticos y el hábito, hábitat y ecosistema de las plantas. Se realizaron análisis estadísticos como Análisis Exploratorios de Datos y Métodos Estadísticos Multivariantes. La longitud y la anchura de los estomas mostraron una muy fuerte correlación, sugiriendo un control genético endógeno que otorga a estos caracteres un considerable valor diagnóstico. Con respecto a las relaciones entre los caracteres estomáticos y el ambiente, encontramos una relación estadísticamente significativa entre la altitud y el índice estomático. También se incluyen interpretaciones de la significación ecológico- selectiva de un conjunto de caracteres estomáticos en diferentes conjuntos de hábitos, hábitats y ecosistemas.
Subject(s)
Biometry/methods , Ecosystem , Ferns/anatomy & histology , Ferns/classification , Plant Stomata/anatomy & histologyABSTRACT
Presence, abundance and reproductive strategies of ferns in disturbed areas of Sierra Nevada, México. Diverse reproductive strategies shown by ferns and lycophytes allow them to colonize a variety of habitats, particularly after the incidence of natural or anthropogenic disturbances. This study assessed the presence, abundance and reproductive strategies of ferns growing in soils of temperate forests with different levels of disturbance at the Sierra Nevada mountain range in the State of Mexico. Vegetation, soil and environmental variables were recorded in 200m² permanent plots located in four forest stands. One-Way ANOVA and Canonical Correspondence Analysis resulted in the recognition of three landscape types defined by the degree of environmental alteration: low, moderate and severe. Also, from five soil samples collected in each stand, germination of ferns and lycophytes was induced. A positive relationship was found between the alteration degree and Cheilanthes abundance. Under a low landscape alteration regime, species richness is restricted to Cheilanthes bonariensis, C. marginata and Pellaea ternifolia subsp. ternifolia. The soil is a reservoir of spores of Cheilanthes and Pellaea ternifolia subsp. ternifolia because their spores can remain viable for different time intervals. Apogamy is the usual reproductive strategy of Cheilanthes species in disturbed ecosystems, although these species also show sexual reproduction in natural ecosystems with adequate water availability. Apogamy may be related to a shorter generation time in comparison with a low disturbed ecosystem. On the other hand, Pellaea ternifolia subsp. ternifolia only has sexual reproduction. Apogamy might be related to a faster generation in comparison with a low disturbed ecosystem. Rev. Biol. Trop. 59 (1): 417-433. Epub 2011 March 01.
Los helechos y licófitas tienen diversas estrategias reproductivas que las habilitan para invadir ambientes perturbados y áreas sin vegetación. En este trabajo se estudió la presencia, abundancia y estrategias reproductivas de individuos que crecen en suelos de bosques templados, en la Sierra Nevada, Estado de México. Se eligieron cuatro zonas en las que se instaló un cuadrante permanente de 200m² para cada una, con el fin de registrar variables edáficas, ambientales y de vegetación. La relación entre variables se pudo explorar con análisis de varianza y Análisis Canónico de Correspondencia dando como resultado tres paisajes con base en el grado de alteración ambiental: baja, moderada y severa. Fueron recolectadas cinco muestras de suelo en cada zona para inducir la germinación de helechos y licofitas, obteniendo resultados que revelan una relación positiva entre el grado de alteración y la abundancia de Cheilanthes. En paisajes con baja alteración, la riqueza de especies fue mayor comparada con paisajes de alteración severa, en la que la presencia de especies se redujo a Cheilanthes bonariensis, C. marginata y Pellaea ternifolia subsp. ternifolia. El suelo fue un banco de esporas de Cheilanthes y Pellaea ternifolia subsp. ternifolia ya que las esporas permanecieron viables por diferentes intervalos de tiempo. La apogamia fue la estrategia reproductiva común para especies de Cheilanthes en ecosistemas alterados, aunque estas especies tienen reproducción sexual en ecosistemas naturales con suficiente agua. Por otro lado, Pellaea ternifolia subsp. ternifolia solo presentó reproducción sexual. La apogamia puede estar relacionada con que la alternancia de generaciones se realice en menos tiempo, comparado con un ecosistema de baja alteración.
Subject(s)
Environmental Monitoring , Ferns/physiology , Trees , Ferns/classification , Mexico , Population Density , Reproduction/physiologyABSTRACT
The extractive exploitation of the tree fern Alsophila setosa Kaulf. alters forest formations and diminishes the availability of micro-habitat for epiphytes. A survey of epiphytic fern communities on A. setosa at 16 study sites in different forest formations in the State of Rio Grande do Sul, Brazil, documented the occurrence of 31 species representing 16 genera and six families. The greatest richness of species occurred in Polypodiaceae (39 percent) and in the genus Asplenium L. (22 percent). Habitual holoepiphyte was the predominant ecological category, representing 61 percent of the species. Similarity analysis demonstrated heterogeneity in the composition of the epiphytic ferns at the study sites and indicated that the vegetation type is not the main factor for floristic difference. The lowest total specific richness (5) was recorded for the seasonal deciduous forest site. The majority of the sites presented similar averages of phorophyte height and epiphyte richness per caudex. In 25 percent of the sites the height of the host plants presented significant correlation with specific richness. Considering that the majority of the epiphytes are habitual and that some of them occur exclusively or preferentially on tree ferns, the maintenance of these host plants in the vegetation is essential for the conservation of epiphytic species.
A exploração extrativista da samambaia arborescente Alsophila setosa Kaulf. altera as formações florestais e diminui a disponibilidade de microhabitats para epífitos. Um levantamento das comunidades de filicíneas epifíticas sobre A. setosa em 16 sítios de diferentes formações florestais, no Estado do Rio Grande do Sul, Brasil, documentou a ocorrência de 31 espécies representando 16 gêneros e seis famílias. A maior riqueza de espécies ocorreu em Polypodiaceae (39 por cento) e no gênero Asplenium L. (22 por cento). A categoria ecológica predominante foi a de holoepífito habitual, representando 61 por cento das espécies. A análise de similaridade evidenciou heterogeneidade na composição de filicíneas epifíticas dos sítios estudados e indicou que o tipo vegetacional não é o principal fator de diferenciação florística. A menor riqueza específica total (5) foi registrada em sítio de floresta estacional decidual. A maioria dos sítios apresentou médias similares de altura dos forófitos e de riqueza de epífitos por cáudice. Em 25 por cento dos sítios, a altura das plantas hospedeiras apresentou correlação significativa com riqueza específica. Considerando que a maioria dos epífitos é habitual e que alguns deles ocorrem exclusivamente ou preferencialmente sobre samambaias arborescentes, a manutenção dessas plantas hospedeiras, na vegetação, é essencial para a conservação de espécies epifíticas.
Subject(s)
Biodiversity , Ferns/classification , Symbiosis , Trees , Brazil , Population DensityABSTRACT
Pteridophytes that indicate environmental alteration in the temperate forest of San Jerónimo Amanalco, Texcoco, México. Pteridophytes that indicate environmental alteration in the San Jerónimo Amanalco temperate forest, Texcoco, Mexico. The patterns of distribution of 26 pteridophyte species were studied as possible indicators of environmental alteration in the temperate forest of San Jerónimo Amanalco, Texcoco, State of Mexico. The presence and abundance of the pteridoflora was studied in relation to edaphic, topographic and vegetation variables in 100 sampling locations within an area of 494 hectares. The relationship between these variables was studied using Canonical Correspondence Analysis. Five landscapes were recognized in the study zone according to the degree of deterioration: severe erosion, erosion, mountain with moderate reversible deterioration, mountain with no evident deterioration, and canyon with no evident deterioration. Cheilanthes bonariensis and Pellaea ternifolia are indicators of environmental degradation. The taxa that only grow in landscapes without apparent alteration are Adiantum andicola, Adiantum poiretii, Argyrochosma incana, Asplenium blepharophorum, Dryopteris pseudo filix-mas, Equisetum hyemale and Pteris cretica. Rev. Biol. Trop. 56 (2): 641-656. Epub 2008 June 30.
Se estudiaron los patrones de distribución de 26 especies de pteridofitas como posibles indicadoras de alteración ambiental en los bosques templados de San Jerónimo Amanalco, Texcoco, Estado de México. La presencia y abundancia de la pteridoflora se relacionó con variables edáficas, topográficas y de vegetación en 100 sitios de muestreo en un área de 494 ha. Con base en el Análisis Canónico de Correspondencia se pudo explorar la relación entre variables. Se determinaron cinco paisajes en el área de estudio con base al grado de deterioro: erosión severa, erosión, montaña con deterioro moderado reversible, montaña sin deterioro aparente y cañada sin deterioro aparente. Cheilanthes bonariensis y Pellaea ternifolia son indicadores de deterioro ambiental. Los táxones que únicamente crecen en paisajes sin alteración aparente son: Adiantum andicola, Adiantum poiretii, Argyrochosma incana, Asplenium blepharophorum, Dryopteris pseudo filix-mas, Equisetum hyemale y Pteris cretica.
Subject(s)
Environmental Monitoring , Ferns/classification , Trees , Ferns/physiology , Mexico , Population DensityABSTRACT
El estudio se realizó en el sendero La Esperanza del parque nacional La Tigra, de septiembre a diciembre de 2002 y de febrero a marzo de 2003. El sendero La Esperanza tiene una longitud de 2.5 km; está ubicado entre 1,940 y 2,120 msnm y se caracteriza por tener una vegetación latifoliada muy diversa. El objetivo principal del estudio fue conocer la diversidad de helechos presentes en el sendero, lo mismo que la etnobotánica de los helechos en Honduras. Se encontró una alta diversidad de helechos (54 especies) dentro de los cuales sobresale Elaphoglossum latifolium, Phlebodium pseudoaureum (helechos epífitos), Alsophila salvinii, Cyathea divergens var. tuerckheimii, Cyathea valdecrenata, Dicksonia gigantea y Lophosoria quadripinnata var. quadripinnata (helechos arborescentes) que son las especies que son comúnmente utilizadas en Honduras. P. pseudoaureum es una de las especies más ampliamente utilizada en Centro y Suramérica. Los helechos arborescentes citados anteriormente son objeto de comercio ilegal en Honduras, Centro América y Sur América, por lo que se encuentran en el Apéndice II de CITES.
Subject(s)
Ferns/classification , Plants, Edible , Plants, Medicinal , Ethnobotany , HondurasABSTRACT
Danaea tuomistoana A. Rojas (Marattiaceae) is described and illustrated as a new species endemic to Costa Rica; it differs from D. crispa Endres et Richb. by a truncate blade, and by the pinnae, which are 2/5-1/2 incised, cuneated at the base and with a flat margin. The range of D. erecta H. Tuomisto & R.C. Moran in Costa Rica is expanded with a new locality in Puntarenas (8 degrees 57'15[quot ] N-82 degrees 50'1[quot ] W, 1,500-1,580 masl).
Se describe e ilustra una especie nueva endémica de helecho de Costa Rica, Danaea tuomistoana A. Rojas (Marattiaceae), caracterizada porque las pinnas son incisas 2/5-1/2 entre la costa y el margen, y son cuneadas en la base. Se amplia el ámbito geográfico de D. erecta H. Tuomisto & R.C. Moran a Puntarenas, Costa Rica.