Thelypteris Research Articles

A Global Phylogenomic Study of the Thelypteridaceae

Abstract— The generic classification of the Thelypteridaceae has been the subject of much controversy. Proposed taxonomic systems have varied from recognizing the approximately 1200 species in the family within the single genus Thelypteris, to systems favoring upwards of 30 genera. Insights on intrafamilial relationships, especially for neotropical taxa, have been gained from recent phylogenetic studies; however, in the most recent classification, 10 of 30 recognized genera are either non-monophyletic or untested. We sequenced 407 nuclear loci for 621 samples, representing all recognized genera and approximately half the known species diversity. These were analyzed using both maximum likelihood analysis of a concatenated matrix and multi-species coalescent methods. Our phylogenomic results, informed by recently published morphological evidence, provide the foundation for a generic classification which recircumscribed 14 genera and recognized seven new genera. The 37 monophyletic genera sampled demonstrate greater geographic coherence than previous taxonomic concepts suggested. Additionally, our results demonstrate that certain morphological characters, such as frond division, are evolutionarily labile and are thus inadequate for defining genera.

Changes in vegetation and environment in the Betula microphylla wetland of Ebinur Lake in Xinjiang, China since 3900 cal. aBP.

Based on the high resolution record of fossil pollen data from a 190 cm sediment profile collected in a Betula microphylla-dominated wetland of Ebinur Lake in Xinjiang, we aimed to reveal the changes of vegetation and environment in this wetland over the past 3900 years. Results showed that, in the period between 3420 cal. aBP and 3940 cal. aBP, this area was a desert landscape dominated by plants from Chenopodiaceae and Artemisia. During the period of 2830-3420 cal. aBP, this area became a desert steppe dominated by Thelypteris palustris, surrounded by desert plants mainly composed of Chenopodiaceae and Artemisia. In the period between 2640 and 2830 cal. aBP, Typha pollen increased and occupied a dominant position. The area thus became a typical desert intrazonal wetland mainly composed of Typha. In the period between 1200 and 2640 cal. aBP, Phragmites australis began to grow vigorously and became the dominant species. In the period between 660 and 1200 cal. aBP, the population of Betula expanded rapidly, and P. australis plants continued to grow. During the period of 340-660 cal. aBP, the population of B. microphylla became the dominant species, and the abundance of P. australis decreased. However, there was still some P. australis wetlands in the area. Over the past 340 years, as the area of wetlands decreased and the surface water gradually declined, B. microphylla had grown on swampy meadow soils with suitable moisture conditions. In the past 200 years especially, the sharp increases of Chenopodiaceae pollen in palynological assemblage was closely related to the enhancement of human activities and climate change, resulting in the degradation of wetlands and endangerment of B. microphylla. Therefore, it is necessary to strengthen the protection of wetlands and endangered species in this area.

Meiosis of Dihaploid <i>Thelypteris decursive-pinnata</i> Produced Artificially by Induced Apogamy

Meiosis of Dihaploid <i>Thelypteris decursive-pinnata</i> Produced Artificially by Induced Apogamy

PLANT-INSECT INTERACTIONS ON DICOTS AND FERNS FROM THE MIOCENE OF ARGENTINA

Fil: Robledo, Juan Manuel. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Centro Cientifico Tecnologico Conicet - Nordeste. Centro de Ecologia Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecologia Aplicada del Litoral; Argentina

The enigma of the Diporotheca palynomorph

In the Quaternary palynological literature, the name Diporotheca rhizophila has come to be applied for fungal spores labelled in the Hugo de Vries-Laboratory (HdV) in Amsterdam (The Netherlands) as Type HdV-143. The widespread occurrence of this taxon in palynological preparations was difficult to understand as the species is definitely known only from Solanum species in Washington State in the USA. Comparison of spores found in palaeoecological and forensic samples with those of the original material (holotype) of D. rhizophila established that the name had been misapplied. Type HdV-143 is distinguished from the holotype by the shape, size, and ornamentation of the ascospores. The spores in the palaeoecological material are navicular to fusiform, and not oval when mature. They develop robust, dark brown, anastomosed ridges from an early stage, and are much longer, measuring 44–59μm and not 25–37μm in length. Similarities in spore structure and development indicate that both can be accommodated in the same genus, but as different species. The specimens from palynological preparations are described here as D. webbiae sp. nov. Circumstantial evidence, gained from palaeoecological analysis, suggests that the new species may be associated with the fern genus Thelypteris, and today occurs most commonly in wet Alnus carr. Similar spores have been reported from the pre-Quaternary fossil record under the generic name Striadiporites.

A diverse fern flora including macrofossils with in situ spores from the late Eocene of southern New Zealand

Three types of fossil fern fronds bearing sporangia with in situ spores are described from the late Eocene Pikopiko Fossil Forest, southern New Zealand. They are referable to the extant fern genus Thelypteris subgenus Cyclosorus (Thelypteridaceae) and an extinct fern of uncertain affinity. The matching of Thelypteris (Cyclosorus) macrofossils to the spore form taxon Laevigatosporites ovatus is recognised for the first time. Five additional fern taxa, including another probable Blechnum, are distinguished on the basis of sterile foliage. The Pikopiko site is one of the richest known Cenozoic fern floras globally and the diversity and abundance of ferns (eight fern macrofossil parataxa and 20 miospore types) implies that ferns dominated the evergreen, tall forest understorey as in modern New Zealand rainforests. At least 40% of modern New Zealand fern families are represented in this 35Ma flora and the discovery of fronds with in situ spores of a type recorded throughout the Cenozoic in New Zealand supports a long history for some fern genera in the region. The abundance of ferns, epiphyllous fungi on many leaves and the presence of palms is evidence for warm humid conditions in Late Eocene New Zealand.

The genus Thelypteris (Thelypteridaceae, Polypodiopsida) in the state of Mato Grosso, Brazil - II - subgenera Amauropelta (Kunze) A.R. Sm., Cyclosorus (Link) C.V. Morton and Steiropteris (C. Chr.) K. Iwats

The family Thelypteridaceae presents considerable species richness throughout the hillside forests and coastal plains of Brazil, as well as in the forests of its southern plateau. This study was based on material collected in 24 trips to various regions within the state of Mato Grosso, as well as material deposited in the principal Brazilian herbaria and in other herbaria. The genus Thelypteris, represented in this study by the subgenera Amauropelta (Kunze) A.R. Sm., Cyclosorus (Link) Morton and Steiropteris (C. Chr.) K. Iwats., comprises 11 species in Mato Grosso and surrounding regions. Keys for identification, descriptions and references to illustrations of diagnostic characters are presented.

Pteridófitas ocorrentes em três fragmentos florestais de um brejo de altitude (Bonito, Pernambuco, Brasil)

O presente trabalho trata do levantamento da flora pteridofítica da Mata da Colônia, Mata da Chuva e Mata da Reserva, no município de Bonito (Pernambuco, Brasil). O local é um brejo de altitude (ca. 700-800m), cuja vegetação é diferenciada da Caatinga típica da região, condicionada principalmente pela altitude elevada, posição geográfica e aspectos climáticos que favorecem o desenvolvimento das pteridófitas. Foram realizadas 17 excursões ao local com o objetivo de coleta e observação das espécies de pteridófitas, sendo acrescentadas aos acervos de herbários, principalmente regionais, cerca de 400 espécimes. O estudo indicou a ocorrência de 93 espécies e duas variedades, distribuídas em 42 gêneros e 17 famílias. As famílias mais representativas foram Polypodiaceae, com 17 espécies e uma variedade, Thelypteridaceae e Pteridaceae, ambas com 11 espécies. O gênero Thelypteris Schmidel apresentou o maior número de espécies, com nove (e uma variedade), seguido por Asplenium L., com sete. A maioria das espécies apresentou-se como herbáceas, terrícolas e hemicriptófitas, ocorrendo no interior da mata. A flora pteridofítica local apresentou riqueza expressiva, trazendo o registro de 12 novas referências para o Estado, das quais sete ainda não haviam sido citadas para a região Nordeste.

Spore morphology of pteridophytes from China IV. Thelypteridaceae 2

The spore morphology of 50 species of 14 genera of Thelypteridaceae from China was investigated under scanning electron microscope (SEM). The spores are monolete and bilaterally symmetric, elliptic, broadly elliptic or subrounded in polar view and kidney-shaped or rounded in equatorial view. The polar axes are 16.8-40.5 #mu#m long, and the equatorial axes are 12.8-58.0 #mu#m. On the basis of their ornamentation pattern, the spores are divided into seven types. (1) Psilate. Spores of Phegopteris decursive-pinnata belong to this type. (2) Retate. Spores of Pseudophegopteris belong to this type. (3) Circinate. Spores of Glaphyropteridopsis, Macrothelypteris oligophlebia belong to this type. (4) Fimbriate-alate. Spores of Parathelypteris subimmersa, P. japonica, P. angulariloba, Cyclogramma auriculata, most species of Macrothelypteris, and some species of Cyclosorus belong to this type. (5) Perforate-lophate. Spores of Parathelypteris, Mesopteris, Metathelypteris and P. connectilis belong to this type. (6) Cristate. Spores of Cyclosorus belong to this type. (7) Echinate. Spores of Leptogramma, Dictyocline, Thelypteris, Pseudocyclosorus subochthodes, most species of Cyclogramma, and some species of Cyclosorus belong to this type. The relationships of some genera in the Thelypteridaceae are discussed based on spore morphology.

Cariologia de algumas espécies de Pteridófitas ocorrentes no nordeste do Brasil

Foram analisadas citogeneticamente, através de técnicas de coloração convencional, sete espécies de pteridófitas homosporadas e três heterosporadas, pertencentes a oito famílias, sendo essas as primeiras contagens cromossômicas para essas espécies no Brasil. Dois parâmetros principais foram analisados no presente estudo: número cromossômico e tipos de núcleos interfásicos. Em algumas espécies, a morfologia cromossômica e a presença de satélites também foram avaliadas. De todas as espécies analisadas, duas não se tem conhecimento de contagens anteriores: Selaginella convoluta (Arn.) Spring (2n=24) e Thelypteris serrata (Cav.) Alston(2n=72), as quais apresentaram números já esperados para os gêneros. Nas demais espécies os números cromossômicos observados foram: Acrostichum aureum L., 2n=60; A. danaeifolium Langsd. & Fisch., 2n=60; Blechnum occidentale L., 2n=124; Cyathea microdonta (Desv.) Domin, 2n=138; Hemionitis palmata L., 2n=60; Marsilea quadrifolia L., 2n=40; Osmunda cinnamomea L., n=22 e Selaginella kraussiana (Kunze) A. Braun., 2n=20. A única alteração cariológica observada foi a presença de um tetravalente no único indivíduo analisado de O. cinnamomea.A estrutura do núcleo interfásico foi mais densamente reticulada, e mais homogeneamente corada nas espécies homosporadas, enquanto nas heterosporadas essa estrutura foi mais variável, apresentando quase sempre cromômeros ou cromocentros em número variado.

Esalquia centennaria, a new genus and new species of Eriophyidae (Acari) from ferns in Brazil

A new genus, Esalquia, is erected to acommodate a new species, Esalquia centennaria (Acari: Eriophyidae), from ferns (Thelypteris sp.) in Brazil.

Levantamento preliminar de Pteridophyta do Parque Estadual do Rio Doce (MG)

Foi realizado um levantamento preliminar de Pteridophyta no Parque Estadual do Rio Doce, sendo coletadas 15 famílias, 27 gêneros c 57 espécies. As famílias encontradas foram Aspleniaceae, Blechnaceae, Dennstaedtiaceae, Dryopteridaceae, Gleicheniaceae, Hymenophyllaceae, Lomariopsidaceae, Osmundaceae, Polypodiaceae, Pteridaceae, Schizaeaceae, Selaginellaceae, Tectariaceae, Thelypteridaceae e Woodsiaceae. Destas, as mais representativas foram Pteridaceae, com 29,8% das espécies distribuídas em quatro gêneros (Adiantum, Hermionitis, Pityrogramma e Pteris) e Thelypteridaceae que apresentou 17,5% das espécies, incluídas em dois gêneros (Macrothelypteris e Thelypteris). As demais famílias variaram de 1,8-8,8% do total das espécies. Das 57 espécies registradas no Parque, 22 apresentaram distribuição bastante restrita, sendo Pteridaceae e Thelypteridaceae as duas famílias de ocorrência mais ampla. Análises do solo das trilhas do Parque revelaram solo ácido, com pH variando de 4,3-5,1, preferencialmente argiloso a muito argiloso e com teores médio a baixo de P, Ca, Al e Mg.

Nuevas citas de Thelypteris subg. Amauropelta (Thelypteridaceae, Pteridophyta) para la flora Argentina

The presence of Thelypteris regnelliana (C. Chr.) Ponce and T. sanctae-catharinae (Rosenst.) Ponce, from NE of Misiones, Argentina, is reported for the first time. Morphological differences with its related taxa, geographic range and ecological characteristics are also given.

New Species of Thelypteris from Puerto Rico

During the course of field work in Puerto Rico since July 1983, many new fern records have been discovered, some of them representing entities new to science. Preliminary to publication of a book about Puerto Rican ferns, a series of short papers has been initiated in order to describe new taxa and discuss other relevant matters. A previous paper (Proctor, 1984) described a new species of Trichomanes; the present one concerns the genus Thelypteris. All the newly discovered Puerto Rican thelypterids belong to subgenera Amauropelta and Goniopteris. In considering Amauropelta, the sectional assignments follow those of Smith (1974). There is no satisfactory modern classification of Goniopteris; therefore the species of this group are not assigned to sections in the present paper. In citing my own collections other than types, the first set of specimens is deposited in the herbarium of the Department of Natural Resources, San Juan, Puerto Rico (SJ); the second and third sets are deposited at the United States National Herbarium (US) and the Institute of Jamaica (IJ), respectively. More than 100 species of Thelypteris occur in the Antilles, and nearly half of these belong to subgenus Amauropelta. The importance of these islands as centers of speciation for the latter taxon is emphasized by the recent discovery of three new Puerto Rican species, now being described. Two of these belong in section Amauropelta, and thus are allied to the species bearing sessile, reddishresinous glands characterized many years ago by Morton (1963).

Nomenclatural Notes on Micronesian Ferns

Since we accept the genus Thelypteris Schmidel in a broad sense, including such segregates as Christella Leveille, it is necessary to provide the appropriate combination for C. guamensis in Thelypteris. Wagner and Grether (Occ. Pap. Bishop Mus. 19:56. 1948) list as Cyclosorus dentatus (Forsk.) Ching the two collections cited by Holttum (Grether 4384, 4385). We could not match these exactly in the considerable series of specimens from Guam labelled T. dentata or T. parasitica. The thickened, reddish, glandular hairs scattered over the underside of the lamina, the pubescence of two ranges of length, the broad fronds with the lowest pinnae only slightly or not reduced, and the short-creeping rhizome would seem to place T. guamensis between T. dentata and T. parasitica. We follow Holttum in recognizing T. guamensis as a separate species, but with some misgivings. The Guam plants of this affinity do not sort well into three, and even less well into two, populations.

Studies on the cytology of South Indian Aspidiaceae.

Cytological investigation on 46 taxa belonging to 43 species under 14 genera of South Indian Aspidiaceae has been carried out. Chromosomal variations such as hybridity, aneuploidy, polyploidy and supernumerary division following meiosis have been encountered, The occurrence of species complexes has been detected in Arachnoides aristata and Cyclosorus interruptus. Bolbitis quoyana, Diplazium silvaticum, Tectaria paleocnemoides and T. decurrens var. minor have been found to be natural hybrids and the possible origin of the former three species has been suggested. Fifty per cent of the total 46 taxa investigated was found to be polyploids. Aneuploid changes in chromosome number have been detected in Thelypteris and Dryopteris. The origin of the different basic numbers in the genus Thelypteris has been discussed. It has been inferred from the present investigation that cytological changes like hybridity, polyploidy and aneuploidy have played an important role in the evolution of the Aspidiaceous ferns in South India.

Preliminary Systematic Studies of the Oak Ferns: Chromatography and Electrophoresis

The Oak Ferns, represented in the United States by Gymnocarpium dryopteris, G. robertianum, and their putative hybrid, have been placed in many different genera. Some authors have placed them in the genus Dryopteris, primarily on the basis of similarities in rachis characters and rhizome scaliness. Others have included them with the Beech Ferns (Phegopteris). The Beech Ferns and the Oak Ferns have in common an elongate creeping rhizome and exindusiate sori (Morton, 1950). Slosson placed them both in the cosmopolitan genus Thelypteris, along with the Marsh Ferns (Wherry, 1961, p. 64). Ching (1933) proposed that the Oak Ferns be regarded as a genus distinct from Phegopteris, and he redefined Gymnocarpium. Currently most workers appear to favor this interpretation; however, other combinations are frequently used, particularly in local and regional floras. Cytological studies have shown a similarity in chromosome size between the Oak Ferns and the Beech Ferns; however, the number differs. The base number of 40 is found in the Oak Ferns (Wagner, 1966). The Broad Beech Fern and the Long Beech Fern have numbers of 30 and 90, respectively (Britton, 1965; Manton, 1950, p. 184). True Dryopteris has a base number of 41 chromosomes (Walker, 1961; 1962). These chromosomes tend to be somewhat larger than those of either the Beech or the Oak Ferns. Chromosome numbers vary within the genus Thelypteris; however, none reported correspond with that of the Oak Ferns. In recent years paper chromatography has gained acceptance as a taxonomic method. Most of the studies have involved taxa at or below the species level. The works of Smith and Levin (1963), Alston and Turner (1963), and others have indicated chromatography to be of particular value in the confirmation of interspecific hybridization. Works with higher taxa have concerned primarily the distribution and occurrence of specific compounds (Kupchan et al. 1961; Mabry et al. 1963). Cellulose acetate electrophoresis has been used in animal taxonomy (Leviton et al. 1964). No reports were located of comparable comparative studies in plants, though immunoelectrophoresis studies have been done with various plant taxa (Gell et al. 1956). This study was undertaken to add further information concerning the status of the Oak Ferns by the consideration of biochemical characteristics, and to explore the value of paper chromatography and cellulose acetate electrophoresis in generic problems. Chromatographs and electrophoretographs of extracts from the Oak Ferns were compared with those of representatives of the Beech Ferns, the Marsh Ferns (Thelypteris), and the Wood Ferns (Dryopteris).

New Species of Thelypteridaceae from India, Ceylon and Burma

The object of the present communication is to describe two species of ferns of the family Thelypteridaceae discovered by Dr. Chandra during field work in India, and also two species represented by specimens noted by me in the Kew Herbarium. This family, comprising in all about one thousand species, almost all tropical, presents many complex problems both of taxonomy and nomenclature. I have been studying the Old World species for the past three years with a view to writing an account of the family in Flora Malesiana. I hope to present a new conspectus of genera, mainly Malesian, at a later date. I must however present a brief explanation of the generic names here adopted. It is generally admitted that the treatment of this family by Copeland in his Genera Filicum (1947) is unsatisfactory. He did not distinguish genera clearly and included them, with other groups not nearly related to them, in his family Aspidiaceae. The fact that natural species-groups within Thelypteridaceae have not yet been clearly distinguished has led various authors to conclude that the only satisfactory arrangement is to include all species in one genus Thelypteris. I believe however that it is possible, though difficult, to recognize species-groups, and that some of them should have generic rank; also that some generic names adopted by Copeland should be used with different circumscriptions. Copeland used the generic name Cyclosorus Link to include a very wide range of species. But that generic name was originally based on one species, C. gongylodes (Schkuhr) Link, a pantropic swamp-fern of similar habit to the temperate Thelypteris palustris Schott. It appears to me that the majority of species of Copeland's Cyclosorus are not at all closely related to the original species, and that they should have other generic names. Sphaerostephanos was established in 1839 by John Smith and applied only to a species with elongate sori; but it seems to me very clear that the original species is closely related to many others in Malesia which have round sori, and as it is the oldest name available for them I am adopting it with a wider circumscription. Here it is used for an Indian species which has hitherto never been fully described; Bory gave it a name in 1833, but his description was brief and has been misinterpreted. Dr. Chandra re-discovered the species, and by chance I recently discovered Bory's original specimens in the Paris Herbarium.

The Californian Species of Thelypteris

The ferns that have been called Dryopteris in most treatments of ferns in California, as for example those by Maxon, in Abrams, Illustrated Flora of the Pacific States, and by Munz, in his A Manual of Southern California Botany, or, erroneously, Aspidium, as by Jepson, in his Manual of the California Flora, have been shown by various authors to belong to two phyletic lines that are now considered to be rather unrelated; extremists even place them in two families-the Aspidiaceae and the Thelypteridaceae. There do not seem sufficient grounds for regarding these species as belonging to different families or perhaps even subfamilies, but it is evident that two distinct groups have been wrongly united under the name Dryopteris. True Dryopteris is represented in California by the species D. argyeta, D. Filix-mas, and D. austriaca.1 The other species-those called by Maxon D. Feei, D. oreopteris, and D. oregana belong to the genus Thelypteris.2 In an article that I published some time ago in the American Fern Journal3 I pointed out some of the distinguishing characters. I reproduce below the key that was given at that time:

Cultivation of American Ferns in England

The accompanying photograph of the northern beechfern, Thelypteris Phegopteris (L.) Slosson, was made by the junior author from fresh material collected near Yonkers, New York. It shows an enlargement of the dorsal view of the middle of a young tinna. The conspicuous hairs along the midrib show up very well, as do those on the leaf-margins also. The veins, which reach the margin, are forked in this upper part of the frond, with the sorus placed dorsally on the distal fork. These are all characters that ally the beech-fern with the genus Thelypteris. An interesting feature of the beech-fern that is not mentioned apparently in the American literature is the presence of two hairs on the walls of the sporangia; these are of two types-acicular and capitate-glandular -and are well described and illustrated' by that careful fern student Christian Luerssen. He indicates that there is some variation in these hairs-sometimes the