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| Publications of Kathleen M. Pryer :chronological alphabetical combined listing:%% Books @book{fds16774, Author = {Argus, G.W. and K.M. Pryer}, Title = {Rare vascular plants in Canada - our natural heritage}, Pages = {277}, Publisher = {Canadian Museum of Nature}, Year = {1990}, Key = {fds16774} } @book{fds353312, Author = {Argus, GW and Pryer, KM}, Title = {Rare vascular plants in Canada; our natural heritage}, Publisher = {Ottawa, Ontario (Canada) Canadian Museum of Nature, Botany Division}, Year = {1990}, url = {http://dx.doi.org/10.5962/bhl.title.132392}, Doi = {10.5962/bhl.title.132392}, Key = {fds353312} } @book{fds353570, Author = {Argus, GW}, Title = {Atlas of the rare vascular plants of Ontario}, Publisher = {Botany Division, National Museum of Natural Sciences = Division de la botanique, Musée national des sciences naturelles}, Year = {1982}, ISBN = {0662504151}, url = {http://dx.doi.org/10.5962/bhl.title.51429}, Doi = {10.5962/bhl.title.51429}, Key = {fds353570} } %% Papers Published @article{fds373350, Author = {Hay, NM and Windham, MD and Mandáková, T and Lysak, MA and Hendriks, KP and Mummenhoff, K and Lens, F and Pryer, KM and Bailey, CD}, Title = {A Hyb-Seq phylogeny of Boechera and related genera using a combination of Angiosperms353 and Brassicaceae-specific bait sets.}, Journal = {American journal of botany}, Volume = {110}, Number = {10}, Pages = {e16226}, Year = {2023}, Month = {October}, url = {http://dx.doi.org/10.1002/ajb2.16226}, Abstract = {<h4>Premise</h4>Although Boechera (Boechereae, Brassicaceae) has become a plant model system for both ecological genomics and evolutionary biology, all previous phylogenetic studies have had limited success in resolving species relationships within the genus. The recent effective application of sequence data from target enrichment approaches to resolve the evolutionary relationships of several other challenging plant groups prompted us to investigate their usefulness in Boechera and Boechereae.<h4>Methods</h4>To resolve the phylogeny of Boechera and closely related genera, we utilized the Hybpiper pipeline to analyze two combined bait sets: Angiosperms353, with broad applicability across flowering plants; and a Brassicaceae-specific bait set designed for use in the mustard family. Relationships for 101 samples representing 81 currently recognized species were inferred from a total of 1114 low-copy nuclear genes using both supermatrix and species coalescence methods.<h4>Results</h4>Our analyses resulted in a well-resolved and highly supported phylogeny of the tribe Boechereae. Boechereae is divided into two major clades, one comprising all western North American species of Boechera, the other encompassing the eight other genera of the tribe. Our understanding of relationships within Boechera is enhanced by the recognition of three core clades that are further subdivided into robust regional species complexes.<h4>Conclusions</h4>This study presents the first broadly sampled, well-resolved phylogeny for most known sexual diploid Boechera. This effort provides the foundation for a new phylogenetically informed taxonomy of Boechera that is crucial for its continued use as a model system.}, Doi = {10.1002/ajb2.16226}, Key = {fds373350} } @article{fds372242, Author = {Hay, NM and Akinwuntan, JV and Cai, V and Windham, MD and Pryer, KM}, Title = {Exploring Past and Future Distributions of the Rare Appalachian Oak Fern Using MaxEnt Modeling}, Journal = {American Fern Journal}, Volume = {113}, Number = {2}, Pages = {109-125}, Year = {2023}, Month = {June}, url = {http://dx.doi.org/10.1640/0002-8444-113.2.109}, Abstract = {Abstract . Anthropogenic climate change is projected to have an especially negative impact on the survival of plants that are dependent on limited microclimatic refugia or that already reside at their climatic extreme. Gymnocarpium appalachianum is a narrowly endemic fern restricted to cold mountaintops and algific vents in the central and southern Appalachian region of eastern North America. It is the much rarer of the two documented diploid parents of the circumboreal allotetraploid G. dryopteris - one of the most widespread fern species on the planet. Gymnocarpium appalachianum is a good case study for forecasting how evolutionarily significant, but rare, species might survive on a warming planet. We utilize an ecological niche modeling approach (MaxEnt) to explore the projected distribution of G. appalachianum under past (Last Glacial Maximum) and future climate models. All known verified herbarium records of G. appalachianum were georeferenced, for a total of 70 occurrence points. Nineteen standard bioclimatic variables extracted from WorldClim were used to model near-current climate projections; representative concentration pathways (RCPs 2.6 and 8.5) were used for future climate projections (2070). The temperature annual range, mean temperature of warmest quarter, precipitation of driest month, precipitation of coldest quarter, and mean diurnal range were identified as the key variables for shaping the distribution of G. appalachianum. An unanticipated result from our analyses is that G. appalachianum has past and current projected habitat suitability in Alaska. Because this overlaps with the current range of G. disjunctum, the other diploid parent of G. dryopteris, it suggests a possible region of origin for this circumboreal tetraploid descendent of G. appalachianum - a research avenue to be pursued in the future. Our study envisions a dire fate for G. appalachianum; its survival will likely require an urgent contingency plan that includes human-mediated population relocation to cooler, northern locations. Understanding the long-term sustainability of narrowly endemic plants such as G. appalachianum is critical in decisions about their management and conservation.}, Doi = {10.1640/0002-8444-113.2.109}, Key = {fds372242} } @article{fds369759, Author = {Windham, MD and Picard, KT and Pryer, KM}, Title = {An in-depth investigation of cryptic taxonomic diversity in the rare endemic mustard Draba maguirei.}, Journal = {American journal of botany}, Volume = {110}, Number = {3}, Pages = {1-22}, Year = {2023}, Month = {March}, url = {http://dx.doi.org/10.1002/ajb2.16138}, Abstract = {<h4>Premise</h4>Previously published evidence suggests that Draba maguirei, a mustard endemic to a few localities in the Bear River, Wellsville, and Wasatch Mountains of northern Utah, may represent a cryptic species complex rather than a single species. Conservation concerns prompted an in-depth systematic study of this taxon and its putative relatives.<h4>Methods</h4>Sampling most known populations of D. maguirei s.l. (D. maguirei var. maguirei and D. maguirei var. burkei), we integrate data from geography, ecology, morphology, cytogenetics and pollen, enzyme electrophoresis, and the phylogenetic analysis of nuclear internal transcribed spacer sequences to explore potential taxonomic diversity in the species complex.<h4>Results</h4>Draba maguirei var. burkei is shown here to be a distinct species (D. burkei) most closely related to D. globosa, rather than to D. maguirei. Within D. maguirei s.s., the northern (high elevation) and southern (low elevation) population clusters are genetically isolated and morphologically distinguishable, leading to the recognition here of the southern taxon as D. maguirei subsp. stonei.<h4>Conclusions</h4>Our study reveals that plants traditionally assigned to D. maguirei comprise three genetically divergent lineages (D. burkei and two newly recognized subspecies of D. maguirei), each exhibiting a different chromosome number and occupying a discrete portion of the geographic range. Although previously overlooked and underappreciated taxonomically, the three taxa are morphologically recognizable based on the distribution and types of trichomes present on the leaves, stems, and fruit. Our clarification of the diversity and distribution of these taxa provides an improved framework for conservation efforts.}, Doi = {10.1002/ajb2.16138}, Key = {fds369759} } @article{fds369760, Author = {Windham, MD and Picard, KT and Pryer, KM}, Title = {Myriopteris grusziae: A New Species from Texas and Oklahoma Segregated from the Chihuahuan Desert Taxon M. scabra (Pteridaceae)}, Journal = {Systematic botany}, Volume = {47}, Number = {3}, Pages = {876-886}, Publisher = {American Society of Plant Taxonomists}, Year = {2022}, Month = {September}, url = {http://dx.doi.org/10.1600/036364422x16573022073590}, Abstract = {Myriopteris scabra (until recently called Cheilanthes horridula) is a xeric-adapted fern species, endemic to the southwestern United States and northern Mexico. It is one of the most recognizable ferns in North America due to the unusual nature of the indument present on its adaxial leaf surfaces. This consists of rigid, multicellular trichomes with glassy, needle-like apices and compact conical bodies that are partially embedded in the leaf surface to form swollen, pustulate bases. Despite the seemingly distinctive nature of M. scabra, published chromosome counts indicate that collections assigned to this taxon encompass both diploids (n = 29) and tetraploids (n = 58). Here we investigate this case of cryptic diversity by integrating data from cytogenetic and spore analyses, observations of sporophyte morphology, and geographic distributions. Myriopteris scabra s.l. is shown to comprise two genetically disparate, morphologically recognizable taxa that exhibit little or no geographic overlap. The tetraploid taxon is described as a new species, M. grusziae, which completely supplants diploid M. scabra in the northeastern portion of its range (central Texas and south-central Oklahoma). This presumed allotetraploid is most like M. scabra but differs in having ultimate segments with adaxial trichomes that are longer, more flexible, mostly linear, and superficially attached. In addition, tetraploid M. grusziae has larger, more abundant scales that largely conceal the dark, sclerified leaf rachises, and it produces consistently larger spores than diploid M. scabra. We hypothesize that M. grusziae is an allotetraploid hybrid that acquired half of its chromosomes from M. scabra. However, the identity of the other diploid parent has yet to be resolved.}, Doi = {10.1600/036364422x16573022073590}, Key = {fds369760} } @article{fds363346, Author = {Windham, MD and Huiet, L and Metzgar, JS and Ranker, TA and Yatskievych, G and Haufler, CH and Pryer, KM}, Title = {Once more unto the breach, dear friends: Resolving the origins and relationships of the Pellaea wrightiana hybrid complex.}, Journal = {American journal of botany}, Volume = {109}, Number = {5}, Pages = {821-850}, Year = {2022}, Month = {May}, url = {http://dx.doi.org/10.1002/ajb2.1850}, Abstract = {<h4>Premise</h4>The taxonomic status of Wright's cliff brake fern, Pellaea wrightiana, has been in dispute ever since it was first described by Hooker in 1858. Previously published evidence suggested that this "taxon" may represent a polyploid complex rather than a single discrete species, a hypothesis tested here using a multifaceted analytical approach.<h4>Methods</h4>Data derived from cytogenetics, spore analyses, leaf morphometrics, enzyme electrophoresis, and phylogenetic analyses of plastid and nuclear DNA sequences are used to elucidate the origin, relationships, and taxonomic circumscription of P. wrightiana.<h4>Results</h4>Plants traditionally assigned to this taxon represent three distinct polyploids. The most widespread, P. wrightiana, is a fertile allotetraploid that arose through hybridization between two divergent diploid species, P. truncata and P. ternifolia. Sterile triploids commonly identified as P. wrightiana, were found to be backcross hybrids between this fertile tetraploid and diploid P. truncata. Relatively common across Arizona and New Mexico, they are here assigned to P. ×wagneri hyb. nov. In addition, occasional sterile tetraploid plants assigned to P. wrightiana are shown here to be hybrids between the fertile allotetraploid and the tetraploid P. ternifolia subsp. arizonica. These tetraploid hybrids originated independently in two regions of parental sympatry (southern Arizona and west Texas) and are here assigned to P. ×gooddingii hyb. nov.<h4>Conclusions</h4>Weaving together data from a diversity of taxonomic approaches, we show that plants identified as P. wrightiana represent three morphologically distinguishable polyploids that have arisen through repeated hybridization events involving the divergent sexual taxa P. ternifolia and P. truncata.}, Doi = {10.1002/ajb2.1850}, Key = {fds363346} } @article{fds365576, Author = {Windham, MD and Pryer, KM}, Title = {FIRST REPORT OF UTAH FRAGILE FERN (CYSTOPTERIS: CYSTOPTERIDACEAE) FROM NEW MEXICO, AND NEW NOMENCLATURAL COMBINATIONS IN CYSTOPTERIS TENNESEENSIS}, Journal = {Journal of the Botanical Research Institute of Texas}, Volume = {16}, Number = {1}, Year = {2022}, Month = {January}, url = {http://dx.doi.org/10.17348/jbrit.v16.i1.1215}, Abstract = {Two recently analyzed collections of Cystopteris from New Mexico represent the first records of Utah fragile fern for the state. Our morphological reexamination of this taxon reveals a single character (the thickness of lateral cell walls in rhizome and petiole base scales) that consistently distinguishes it from tetraploid Cystopteris tennesseensis. Therefore, we propose the new combination C. tennesseensis subsp. utahensis to accommodate these disjunct, tetraploid populations of the southwestern United States. A second new combination, C. tennesseensis subsp. laurentiana, is provided for morphologically similar hexaploids from the northeastern U.S. and southern Canada.}, Doi = {10.17348/jbrit.v16.i1.1215}, Key = {fds365576} } @article{fds365577, Author = {Windham, MD and Pryer, KM and Allphin, L}, Title = {AN OVERVIEW OF NEW MEXICAN BOECHERA (BRASSICACEAE) INCLUDING THREE NEW SPECIES AND THIRTEEN NEW NOMENCLATURAL COMBINATIONS}, Journal = {Journal of the Botanical Research Institute of Texas}, Volume = {16}, Number = {1}, Pages = {9-24}, Year = {2022}, Month = {January}, url = {http://dx.doi.org/10.17348/jbrit.v16.i1.1216}, Abstract = {To provide a nomenclatural foundation for the treatment of Boechera in the soon-to-be-published Flora of New Mexico, we here discuss the current taxonomic situation in the genus, name three new species (B. austromontana, B. pseudoconsanguinea, and B. quadrangulensis), provide new combinations for B. kelseyana, B. thompsonii, and B. villosa as subspecies of B. crandallii, and propose subspecific names under B. perennans for nine taxa currently treated as species.}, Doi = {10.17348/jbrit.v16.i1.1216}, Key = {fds365577} } @article{fds359722, Author = {Heron, K and Windham, MD and Farrar, DR and Pryer, KM}, Title = {Looking Back on 130 Years of Fern and Lycophyte Research in Glacier National Park, Montana: A Modern Taxonomic Account}, Journal = {American fern journal.}, Volume = {111}, Number = {4}, Pages = {223-250}, Publisher = {American Fern Society}, Year = {2021}, Month = {November}, url = {http://dx.doi.org/10.1640/0002-8444-111.4.223}, Abstract = {Glacier National Park encompasses over one million acres in the mountains of northwestern Montana, along the United States–Canada border. Our survey of online databases indicates that the earliest extant fern and lycophyte collections from this area were taken by Robert S. Williams in 1892. In the summer of 1919, Paul C. Standley, a botanist with the United States National Museum, conducted a survey of the flora of the newly created Park and recorded 39 species of ferns and lycophytes. In 2002, a revised flora for the Park by Peter Lesica increased this number to 61. Here we summarize 130 years of collections-based research on the ferns and lycophytes of Glacier National Park, documenting how our understanding of the flora has changed through time. In the summer of 2019, the lead author conducted a field survey to relocate as many ferns and lycophytes as possible within park boundaries. In parallel, we scoured herbarium online portals and databases for high-resolution digitized specimen images to confirm or refute historical vouchers of ferns and lycophytes collected from the Park. In a few cases, specimen loans were requested from herbaria to confirm our determinations. The results from our combined field and online herbarium studies are presented here. Of the 61 taxa recognized by Lesica in 2002, we were able to confirm all but seven. In sum, we recognize here a total of 71 fern and lycophyte taxa for the Park. Most previously unreported taxa belong to Botrychium, a genus that has seen a flurry of recent taxonomic work by co-author Farrar and collaborators. These new data are presented here together with updated nomenclature and discussion to provide a current taxonomic account of the fourteen fern and lycophyte families known to occur in Glacier National Park. We anticipate this study will provide a useful foundation for further investigations in the Park.}, Doi = {10.1640/0002-8444-111.4.223}, Key = {fds359722} } @article{fds359082, Author = {Sosa, K and Pryer, KM and Huiet, L and Yatskievych, G and Windham, MD}, Title = {Cheilanthes ecuadorensis: A New Species of Cheilanthes s. s. (Pteridaceae) from Northern South America}, Journal = {Systematic botany}, Volume = {46}, Number = {2}, Pages = {249-259}, Year = {2021}, Month = {August}, url = {http://dx.doi.org/10.1600/036364421x16231782047280}, Abstract = {Ongoing research on the taxonomically complex genus Cheilanthes (Pteridaceae; Cheilanthoideae) has resulted in the identification of a new species from Loja Province in Ecuador, Cheilanthes ecuadorensis, described and illustrated herein. Originally collected in 1988 and identified as C. cf. rufopunctata, C. ecuadorensis is clearly distinct from that species in having pubescent adaxial blade surfaces and narrow, poorly-differentiated false indusia (rather than the glabrous adaxial surfaces and wide false indusia of C. rufopunctata). Among the South American species currently included in Cheilanthes, C. ecuadorensis is superficially most similar to C. pilosa. However, our molecular phylogenetic analyses indicate that C. ecuadorensis is sister to C. micropteris, the morphologically disparate generitype of Cheilanthes. Here we examine the phylogenetic relationships, morphology, cytogenetics, and geography of these four South American Cheilanthes species in a study that, once again, highlights the importance of herbaria in the process of new species discovery.}, Doi = {10.1600/036364421x16231782047280}, Key = {fds359082} } @article{fds356426, Author = {Fauskee, BD and Sigel, EM and Pryer, KM and Grusz, AL}, Title = {Variation in frequency of plastid RNA editing within Adiantum implies rapid evolution in fern plastomes.}, Journal = {American journal of botany}, Volume = {108}, Number = {5}, Pages = {820-827}, Year = {2021}, Month = {May}, url = {http://dx.doi.org/10.1002/ajb2.1649}, Abstract = {<h4>Premise</h4>Recent studies of plant RNA editing have demonstrated that the number of editing sites can vary widely among large taxonomic groups (orders, families). Yet, very little is known about intrageneric variation in frequency of plant RNA editing, and no study has been conducted in ferns.<h4>Methods</h4>We determined plastid RNA-editing counts for two species of Adiantum (Pteridaceae), A. shastense and A. aleuticum, by implementing a pipeline that integrated read-mapping and SNP-calling software to identify RNA-editing sites. We then compared the edits found in A. aleuticum and A. shastense with previously published edits from A. capillus-veneris by generating alignments for each plastid gene.<h4>Results</h4>We found direct evidence for 505 plastid RNA-editing sites in A. aleuticum and 509 in A. shastense, compared with 350 sites in A. capillus-veneris. We observed striking variation in the number and location of the RNA-editing sites among the three species, with reverse (U-to-C) editing sites showing a higher degree of conservation than forward (C-to-U) sites. Additionally, sites involving start and stop codons were highly conserved.<h4>Conclusions</h4>Variation in the frequency of RNA editing within Adiantum implies that RNA-editing sites can be rapidly gained or lost throughout evolution. However, varying degrees of conservation between both C-to-U and U-to-C sites and sites in start or stop codons, versus other codons, hints at the likely independent origin of both types of edits and a potential selective advantage conferred by RNA editing.}, Doi = {10.1002/ajb2.1649}, Key = {fds356426} } @article{fds355319, Author = {Grusz, AL and Windham, MD and Picard, KT and Pryer, KM and Schuettpelz, E and Haufler, CH}, Title = {A drought-driven model for the evolution of obligate apomixis in ferns: evidence from pellaeids (Pteridaceae).}, Journal = {American journal of botany}, Volume = {108}, Number = {2}, Pages = {263-283}, Year = {2021}, Month = {February}, url = {http://dx.doi.org/10.1002/ajb2.1611}, Abstract = {<h4>Premise</h4>Xeric environments impose major constraints on the fern life cycle, yet many lineages overcome these limitations by evolving apomixis. Here, we synthesize studies of apomixis in ferns and present an evidence-based model for the evolution and establishment of this reproductive strategy, focusing on genetic and environmental factors associated with its two defining traits: the production of "unreduced" spores (n = 2n) and the initiation of sporophytes from gametophyte tissue (i.e., diplospory and apogamy, respectively).<h4>Methods</h4>We evaluated existing literature in light of the hypothesis that abiotic characteristics of desert environments (e.g., extreme diurnal temperature fluctuations, high light intensity, and water limitation) drive the evolution of obligate apomixis. Pellaeid ferns (Cheilanthoideae: Pteridaceae) were examined in detail, as an illustrative example. We reconstructed a plastid (rbcL, trnG-trnR, atpA) phylogeny for the clade and mapped reproductive mode (sexual versus apomictic) and ploidy across the resulting tree.<h4>Results</h4>Our six-stage model for the evolution of obligate apomixis in ferns emphasizes the role played by drought and associated abiotic conditions in the establishment of this reproductive approach. Furthermore, our updated phylogeny of pellaeid ferns reveals repeated origins of obligate apomixis and shows an increase in the frequency of apomixis, and rarity of sexual reproduction, among taxa inhabiting increasingly dry North American deserts.<h4>Conclusions</h4>Our findings reinforce aspects of other evolutionary, physiological, developmental, and omics-based studies, indicating a strong association between abiotic factors and the establishment of obligate apomixis in ferns. Water limitation, in particular, appears critical to establishment of this reproductive mode.}, Doi = {10.1002/ajb2.1611}, Key = {fds355319} } @article{fds349226, Author = {Kao, T and Rothfels, CJ and Melgoza‐Castillo, A and Pryer, KM and Windham, MD}, Title = {Infraspecific diversification of the star cloak fern (Notholaena standleyi) in the deserts of the United States and Mexico}, Journal = {American Journal of Botany}, Volume = {107}, Number = {4}, Pages = {658-675}, Year = {2020}, url = {http://dx.doi.org/10.1002/ajb2.1461}, Abstract = {PREMISE:Not all ferns grow in moist and shaded habitats. One well-known example is Notholaena standleyi, a species that thrives in deserts of the southwestern United States and Mexico. This species exhibits several "chemotypes" that differ in farina (flavonoid exudates) color and chemistry. By integrating data from molecular phylogenetics, cytology, biochemistry, and biogeography, we circumscribed the major evolutionary lineages within N. standleyi and reconstructed their diversification histories. METHODS:Forty-eight samples were selected from across the geographic distribution of N. standleyi. Phylogenetic relationships were inferred using four plastid and five nuclear markers. Ploidy levels were inferred using spore sizes calibrated by chromosome counts, and farina chemistry was compared using thin-layer chromatography. RESULTS:Four clades are recognized, three of which roughly correspond to previously recognized chemotypes. The diploid clades G and Y are found in the Sonoran and Chihuahuan deserts, respectively; they are estimated to have diverged in the Pleistocene, congruent with the postulated timing of climatological events separating these two deserts. Clade P/YG is tetraploid and partially overlaps the distribution of clade Y in the eastern Chihuahuan Desert. It is apparently confined to limestone, a geologic substrate rarely occupied by members of the other clades. The cryptic (C) clade, a diploid group known only from southern Mexico and highly disjunct from the other three clades, is newly recognized here. CONCLUSIONS:Our results reveal a complex intraspecific diversification history of N. standleyi, traceable to a variety of evolutionary drivers including classic allopatry, parapatry with or without changes in geologic substrate, and sympatric divergence through polyploidization.}, Doi = {10.1002/ajb2.1461}, Key = {fds349226} } @article{fds349481, Author = {Windham, MD and Pryer, KM and Poindexter, DB and Li, F and Rothfels, CJ and Beck, JB}, Title = {A step‐by‐step protocol for meiotic chromosome counts in flowering plants: A powerful and economical technique revisited}, Journal = {Applications in Plant Sciences}, Volume = {8}, Number = {4}, Pages = {e11342}, Publisher = {John Wiley & Sons, Inc.}, Year = {2020}, url = {http://dx.doi.org/10.1002/aps3.11342}, Abstract = {<h4>Premise</h4>Counting chromosomes is a fundamental botanical technique, yet it is often intimidating and increasingly sidestepped. Once mastered, the basic protocol can be applied to a broad range of taxa and research questions. It also reveals an aspect of the plant genome that is accessible with only the most basic of resources-access to a microscope with 1000× magnification is the most limiting factor.<h4>Methods and results</h4>Here we provide a detailed protocol for choosing, staining, and squashing angiosperm pollen mother cells. The protocol is supplemented by figures and two demonstration videos.<h4>Conclusions</h4>The protocol we provide will hopefully demystify and reinvigorate a powerful and once commonplace botanical technique that is available to researchers regardless of their location and resources.}, Doi = {10.1002/aps3.11342}, Key = {fds349481} } @article{fds350496, Author = {Pryer, KM and Tomasi, C and Wang, X and Meineke, EK and Windham, MD}, Title = {Using computer vision on herbarium specimen images to discriminate among closely related horsetails (Equisetum)}, Journal = {Applications in plant sciences}, Volume = {8}, Number = {6}, Pages = {e11372}, Year = {2020}, url = {http://dx.doi.org/10.1002/aps3.11372}, Abstract = {Premise:Equisetum is a distinctive vascular plant genus with 15 extant species worldwide. Species identification is complicated by morphological plasticity and frequent hybridization events, leading to a disproportionately high number of misidentified specimens. These may be correctly identified by applying appropriate computer vision tools. Methods:We hypothesize that aerial stem nodes can provide enough information to distinguish among Equisetum hyemale, E. laevigatum, and E . ×ferrissii, the latter being a hybrid between the other two. An object detector was trained to find nodes on a given image and to distinguish E. hyemale nodes from those of E. laevigatum. A classifier then took statistics from the detection results and classified the given image into one of the three taxa. Both detector and classifier were trained and tested on expert manually annotated images. Results:In our exploratory test set of 30 images, our detector/classifier combination identified all 10 E. laevigatum images correctly, as well as nine out of 10 E. hyemale images, and eight out of 10 E. ×ferrissii images, for a 90% classification accuracy. Discussion:Our results support the notion that computer vision may help with the identification of herbarium specimens once enough manual annotations become available.}, Doi = {10.1002/aps3.11372}, Key = {fds350496} } @article{fds350495, Author = {Meineke, EK and Tomasi, C and Yuan, S and Pryer, KM}, Title = {Applying machine learning to investigate long‐term insect–plant interactions preserved on digitized herbarium specimens}, Journal = {Applications in plant sciences}, Volume = {8}, Number = {6}, Pages = {e11369}, Year = {2020}, url = {http://dx.doi.org/10.1002/aps3.11369}, Abstract = {<h4>Premise</h4>Despite the economic significance of insect damage to plants (i.e., herbivory), long-term data documenting changes in herbivory are limited. Millions of pressed plant specimens are now available online and can be used to collect big data on plant-insect interactions during the Anthropocene.<h4>Methods</h4>We initiated development of machine learning methods to automate extraction of herbivory data from herbarium specimens by training an insect damage detector and a damage type classifier on two distantly related plant species (<i>Quercus bicolor</i> and <i>Onoclea sensibilis</i>). We experimented with (1) classifying six types of herbivory and two control categories of undamaged leaf, and (2) detecting two of the damage categories for which several hundred annotations were available.<h4>Results</h4>Damage detection results were mixed, with a mean average precision of 45% in the simultaneous detection and classification of two types of damage. However, damage classification on hand-drawn boxes identified the correct type of herbivory 81.5% of the time in eight categories. The damage classifier was accurate for categories with 100 or more test samples.<h4>Discussion</h4>These tools are a promising first step for the automation of herbivory data collection. We describe ongoing efforts to increase the accuracy of these models, allowing researchers to extract similar data and apply them to biological hypotheses.}, Doi = {10.1002/aps3.11369}, Key = {fds350495} } @article{fds343683, Author = {Yang, EJ and Yoo, CY and Liu, J and Wang, H and Cao, J and Li, F and Pryer, KM and Sun, T and Weigel, D and Zhou, P}, Title = {NCP activates chloroplast transcription by controlling phytochrome-dependent dual nuclear and plastidial switches}, Journal = {Nature communications}, Volume = {10}, Number = {1}, Pages = {1-13}, Publisher = {Nature Publishing Group}, Year = {2019}, url = {http://dx.doi.org/10.1038/s41467-019-10517-1}, Abstract = {Phytochromes initiate chloroplast biogenesis by activating genes encoding the photosynthetic apparatus, including photosynthesis-associated plastid-encoded genes (PhAPGs). PhAPGs are transcribed by a bacterial-type RNA polymerase (PEP), but how phytochromes in the nucleus activate chloroplast gene expression remains enigmatic. We report here a forward genetic screen in Arabidopsis that identified NUCLEAR CONTROL OF PEP ACTIVITY (NCP) as a necessary component of phytochrome signaling for PhAPG activation. NCP is dual-targeted to plastids and the nucleus. While nuclear NCP mediates the degradation of two repressors of chloroplast biogenesis, PIF1 and PIF3, NCP in plastids promotes the assembly of the PEP complex for PhAPG transcription. NCP and its paralog RCB are non-catalytic thioredoxin-like proteins that diverged in seed plants to adopt nonredundant functions in phytochrome signaling. These results support a model in which phytochromes control PhAPG expression through light-dependent double nuclear and plastidial switches that are linked by evolutionarily conserved and dual-localized regulatory proteins.}, Doi = {10.1038/s41467-019-10517-1}, Key = {fds343683} } @article{fds341364, Author = {Eily, AN and Pryer, KM and Li, F}, Title = {A first glimpse at genes important to the Azolla–Nostoc symbiosis}, Journal = {Symbiosis}, Volume = {78}, Number = {2}, Pages = {149-162}, Publisher = {Springer Netherlands}, Year = {2019}, url = {http://dx.doi.org/10.1007/s13199-019-00599-2}, Abstract = {Azolla is a small genus of diminutive aquatic ferns with a surprisingly vast potential to benefit the environment and agriculture, as well as to provide insight into the evolution of plant-cyanobacterial symbioses. This capability is derived from the unique relationship Azolla spp. have with their obligate, nitrogen-fixing cyanobacterial symbiont, Nostoc azollae, that resides in their leaves. Although previous work has specified the importance of the exchange of ammonium and sucrose metabolites between these two partners, we have yet to determine the underlying molecular mechanisms that make this symbiosis so successful. The newly sequenced and annotated reference genome of Azolla filiculoides has allowed us to investigate gene expression profiles of A. filiculoides—both with and without its obligate cyanobiont, N. azollae—revealing genes potentially essential to the Azolla-Nostoc symbiosis. We observed the absence of differentially expressed glutamine synthetase (GS) and glutamate synthase (GOGAT) genes, leading to questions about how A. filiculoides regulates the machinery it uses for nitrogen assimilation. Ushering A. filiculoides into the era of transcriptomics sets the stage to truly begin to understand the uniqueness of the Azolla-Nostoc symbiosis.}, Doi = {10.1007/s13199-019-00599-2}, Key = {fds341364} } @article{fds343471, Author = {Kao, T and Pryer, KM and Freund, FD and Windham, MD and Rothfels, CJ}, Title = {Low-copy nuclear sequence data confirm complex patterns of farina evolution in notholaenid ferns (Pteridaceae)}, Journal = {Molecular phylogenetics and evolution}, Volume = {138}, Pages = {139-155}, Publisher = {Academic Press}, Year = {2019}, url = {http://dx.doi.org/10.1016/j.ympev.2019.05.016}, Abstract = {Notholaenids are an unusual group of ferns that have adapted to, and diversified within, the deserts of Mexico and the southwestern United States. With approximately 40 species, this group is noted for being desiccation-tolerant and having "farina"-powdery exudates of lipophilic flavonoid aglycones-that occur on both the gametophytic and sporophytic phases of their life cycle. The most recent circumscription of notholaenids based on plastid markers surprisingly suggests that several morphological characters, including the expression of farina, are homoplasious. In a striking case of convergence, Notholaena standleyi appears to be distantly related to core Notholaena, with several taxa not before associated with Notholaena nested between them. Such conflicts can be due to morphological homoplasy resulting from adaptive convergence or, alternatively, the plastid phylogeny itself might be misleading, diverging from the true species tree due to incomplete lineage sorting, hybridization, or other factors. In this study, we present a species phylogeny for notholaenid ferns, using four low-copy nuclear loci and concatenated data from three plastid loci. A total of 61 individuals (49 notholaenids and 12 outgroup taxa) were sampled, including 31 out of 37 recognized notholaenid species. The homeologous/allelic nuclear sequences were retrieved using PacBio sequencing and the PURC bioinformatics pipeline. Each dataset was first analyzed individually using maximum likelihood and Bayesian inference, and the species phylogeny was inferred using *BEAST. Although we observed several incongruences between the nuclear and plastid phylogenies, our principal results are broadly congruent with previous inferences based on plastid data. By mapping the presence of farina and their biochemical constitutions on our consensus phylogenetic tree, we confirmed that the characters are indeed homoplastic and have complex evolutionary histories. Hybridization among recognized species of the notholaenid clade appears to be relatively rare compared to that observed in other well-studied fern genera.}, Doi = {10.1016/j.ympev.2019.05.016}, Key = {fds343471} } @article{fds346477, Author = {George, LO and Pryer, KM and Kao, T and Huiet, L and Windham, MD}, Title = {Baja: A New Monospecific Genus Segregated from Cheilanthes sl (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {44}, Number = {3}, Pages = {471-482}, Publisher = {American Society of Plant Taxonomists}, Year = {2019}, url = {http://dx.doi.org/10.1600/036364419x15620113920536}, Abstract = {The phylogenetic position of Cheilanthes brandegeei, a fern endemic to the Baja California Peninsula of Mexico, was investigated using three plastid markers (atpA, rbcL, trnG-R) and comparative morphology. Here we present robust evidence for the recognition of C. brandegeei as a member of the bommeriids, the sister clade to all other cheilanthoid ferns, and evidence that it is sister to all Bommeria species within that clade. Because of its distinctive morphology within the bommeriid clade (pinnate leaf architecture, well-developed pseudoindusium, and narrow, concolorous red-brown rhizome scales), here we propose the new genus Baja to accommodate it. Our results place Baja brandegeei together with other taxa that have a distribution in the Baja California Peninsula and mainland Mexico, rather than with hypothesized congeners in South America and Africa. Morphological characters traditionally used to classify this species as a Cheilanthes (patterns of sporangial distribution, presence of a well-developed pseudoindusium, and fractiferous petioles) are extensively homoplasious across cheilanthoids. We identify three characters that unite the newly expanded bommeriid clade: leaf indument of acicular trichomes, reticulate-cristate perispore morphology, and lateral initiation of the gametophyte meristem.}, Doi = {10.1600/036364419x15620113920536}, Key = {fds346477} } @article{fds346456, Author = {Sigel, EM and Der, JP and Windham, MD and Pryer, KM}, Title = {Expression Level Dominance and Homeolog Expression Bias in Recurrent Origins of the Allopolyploid Fern Polypodium hesperium}, Journal = {American Fern Journal}, Volume = {109}, Number = {3}, Pages = {224-247}, Publisher = {The American Fern Society}, Year = {2019}, url = {http://dx.doi.org/10.1640/0002-8444-109.3.224}, Abstract = {Allopolyploidization is a common mode of speciation in ferns with many taxa having formed recurrently from distinct hybridization events between the same parent species. Each hybridization event marks the union of divergent parental gene copies, or homeologs, and the formation of an independently derived lineage. Little is known about the effects of recurrent origins on the genomic composition and phenotypic variation of allopolyploid fern taxa. To begin to address this knowledge gap, we investigated gene expression patterns in two naturally formed, independently derived lineages of the allotetraploid fern Polypodium hesperium relative to its diploid progenitor species, Polypodium amorphum and Polypodium glycyrrhiza. Using RNA-sequencing to survey total gene expression levels for 19194 genes and homeolog-specific expression for 1073 genes, we found that, in general, gene expression in both lineages of P. hesperium was biased toward P. amorphum—both by mirroring expression levels of P. amorphum and preferentially expressing homeologs derived from P. amorphum. However, we recovered substantial expression variation between the two lineages at the level of individual genes and among individual specimens. Our results align with similar transcriptome profile studies of angiosperms, suggesting that expression in many allopolyploid plants reflects the dominance of a specific parental subgenome, but that recurrent origins impart substantial expression, or phenotypic, variation to allopolyploid taxa.}, Doi = {10.1640/0002-8444-109.3.224}, Key = {fds346456} } @article{fds330472, Author = {Dijkhuizen, LW and Brouwer, P and Bolhuis, H and Reichart, G and Koppers, N and Huettel, B and Bolger, AM and Li, F and Cheng, S and Liu, X}, Title = {Is there foul play in the leaf pocket? The metagenome of floating fern Azolla reveals endophytes that do not fix N2 but may denitrify}, Journal = {New Phytologist}, Volume = {217}, Number = {1}, Pages = {453-466}, Year = {2018}, url = {http://dx.doi.org/10.1111/nph.14843}, Abstract = {Dinitrogen fixation by Nostoc azollae residing in specialized leaf pockets supports prolific growth of the floating fern Azolla filiculoides. To evaluate contributions by further microorganisms, the A. filiculoides microbiome and nitrogen metabolism in bacteria persistently associated with Azolla ferns were characterized. A metagenomic approach was taken complemented by detection of N2 O released and nitrogen isotope determinations of fern biomass. Ribosomal RNA genes in sequenced DNA of natural ferns, their enriched leaf pockets and water filtrate from the surrounding ditch established that bacteria of A. filiculoides differed entirely from surrounding water and revealed species of the order Rhizobiales. Analyses of seven cultivated Azolla species confirmed persistent association with Rhizobiales. Two distinct nearly full-length Rhizobiales genomes were identified in leaf-pocket-enriched samples from ditch grown A. filiculoides. Their annotation revealed genes for denitrification but not N2 -fixation. 15 N2 incorporation was active in ferns with N. azollae but not in ferns without. N2 O was not detectably released from surface-sterilized ferns with the Rhizobiales. N2 -fixing N. azollae, we conclude, dominated the microbiome of Azolla ferns. The persistent but less abundant heterotrophic Rhizobiales bacteria possibly contributed to lowering O2 levels in leaf pockets but did not release detectable amounts of the strong greenhouse gas N2 O.}, Doi = {10.1111/nph.14843}, Key = {fds330472} } @article{fds336265, Author = {Li, F and Brouwer, P and Carretero-Paulet, L and Cheng, S and De Vries, J and Delaux, P and Eily, A and Koppers, N and Kuo, L and Li, Z}, Title = {Fern genomes elucidate land plant evolution and cyanobacterial symbioses}, Journal = {Nature plants}, Volume = {4}, Number = {7}, Pages = {460-472}, Publisher = {Nature Publishing Group}, Year = {2018}, url = {http://dx.doi.org/10.1038/s41477-018-0188-8}, Abstract = {Ferns are the closest sister group to all seed plants, yet little is known about their genomes other than that they are generally colossal. Here, we report on the genomes of Azolla filiculoides and Salvinia cucullata (Salviniales) and present evidence for episodic whole-genome duplication in ferns-one at the base of 'core leptosporangiates' and one specific to Azolla. One fern-specific gene that we identified, recently shown to confer high insect resistance, seems to have been derived from bacteria through horizontal gene transfer. Azolla coexists in a unique symbiosis with N<sub>2</sub>-fixing cyanobacteria, and we demonstrate a clear pattern of cospeciation between the two partners. Furthermore, the Azolla genome lacks genes that are common to arbuscular mycorrhizal and root nodule symbioses, and we identify several putative transporter genes specific to Azolla-cyanobacterial symbiosis. These genomic resources will help in exploring the biotechnological potential of Azolla and address fundamental questions in the evolution of plant life.}, Doi = {10.1038/s41477-018-0188-8}, Key = {fds336265} } @article{fds336266, Author = {Huiet, L and Li, F and Kao, T and Prado, J and Smith, AR and Schuettpelz, E and Pryeri, KM}, Title = {A worldwide phylogeny of Adiantum (Pteridaceae) reveals remarkable convergent evolution in leaf blade architecture}, Journal = {Taxon}, Volume = {67}, Number = {3}, Pages = {488-502}, Publisher = {WILEY}, Year = {2018}, url = {http://dx.doi.org/10.12705/673.3}, Abstract = {Adiantum is among the most distinctive and easily recognized leptosporangiate fern genera. Despite encompassing an astonishing range of leaf complexity, all species of Adiantum share a unique character state not observed in other ferns: sporangia borne directly on the reflexed leaf margin or “false indusium” (pseudoindusium). The over 200 species of Adiantum span six continents and are nearly all terrestrial. Here, we present one of the most comprehensive phylogenies for any large (200+ spp.) monophyletic, subcosmopolitan genus of ferns to date. We build upon previous datasets, providing new data from four plastid markers (rbcL, atpA, rpoA, chlN) for 146 taxa. All sampled taxa can be unequivocally assigned to one of nine robustly supported clades. Although some of these unite to form larger, well-supported lineages, the backbone of our phylogeny has several short branches and generally weak support, making it difficult to accurately assess deep relationships. Our maximum likelihood-based ancestral character state reconstructions of leaf blade architecture reveal remarkable convergent evolution across multiple clades for nearly all leaf forms. A single unique synapomorphy—leaves once-pinnate, usually with prolonged rooting tips—defines the philippense clade. Although a rare occurrence in Adiantum, simple leaves occur in three distinct clades (davidii, philippense, peruvianum). Most taxa have leaves that are more than once-pinnate, and only a few of these (in the formosum and pedatum clades) exhibit the distinct pseudopedate form. Distributional ranges for each of the terminal taxa show that most species (75%) are restricted to only one of six major biogeographical regions. Forty-eight of our sampled species (nearly one-third) are endemic to South America.}, Doi = {10.12705/673.3}, Key = {fds336266} } @article{fds336267, Author = {Schuettpelz, E and Rouhan, G and Pryer, KM and Rothfels, CJ and Prado, J and Sundue, MA and Windham, MD and Moran, RC and Smith, AR}, Title = {Are there too many fern genera?}, Journal = {Taxon}, Volume = {67}, Number = {3}, Pages = {473-480}, Publisher = {WILEY}, Year = {2018}, url = {http://dx.doi.org/10.12705/673.1}, Doi = {10.12705/673.1}, Key = {fds336267} } @article{fds332387, Author = {Song, M and Kuo, L and Huiet, L and Pryer, KM and Rothfels, CJ and Li, F}, Title = {A novel chloroplast gene reported for flagellate plants}, Journal = {American journal of botany}, Volume = {105}, Number = {1}, Pages = {117-121}, Year = {2018}, url = {http://dx.doi.org/10.1002/ajb2.1010}, Abstract = {PREMISE OF THE STUDY:Gene space in plant plastid genomes is well characterized and annotated, yet we discovered an unrecognized open reading frame (ORF) in the fern lineage that is conserved across flagellate plants. METHODS:We initially detected a putative uncharacterized ORF by the existence of a highly conserved region between rps16 and matK in a series of matK alignments of leptosporangiate ferns. We mined available plastid genomes for this ORF, which we now refer to as ycf94, to infer evolutionary selection pressures and assist in functional prediction. To further examine the transcription of ycf94, we assembled the plastid genome and sequenced the transcriptome of the leptosporangiate fern Adiantum shastense Huiet & A.R. Sm. KEY RESULTS:The ycf94 predicted protein has a distinct transmembrane domain but with no sequence homology to other proteins with known function. The nonsynonymous/synonymous substitution rate ratio of ycf94 is on par with other fern plastid protein-encoding genes, and additional homologs can be found in a few lycophyte, moss, hornwort, and liverwort plastid genomes. Homologs of ycf94 were not found in seed plants. In addition, we report a high level of RNA editing for ycf94 transcripts-a hallmark of protein-coding genes in fern plastomes. CONCLUSIONS:The degree of sequence conservation, together with the presence of a distinct transmembrane domain and RNA-editing sites, suggests that ycf94 is a protein-coding gene of functional significance in ferns and, potentially, bryophytes and lycophytes. However, the origin and exact function of this gene require further investigation.}, Doi = {10.1002/ajb2.1010}, Key = {fds332387} } @article{fds339280, Author = {Sigel, EM and Schuettpelz, E and Pryer, KM and Der, JP}, Title = {Overlapping patterns of gene expression between gametophyte and sporophyte phases in the fern Polypodium amorphum (Polypodiales)}, Journal = {Frontiers in plant science}, Volume = {9}, Pages = {1450}, Publisher = {Frontiers}, Year = {2018}, url = {http://dx.doi.org/10.3389/fpls.2018.01450}, Abstract = {Ferns are unique among land plants in having sporophyte and gametophyte phases that are both free living and fully independent. Here, we examine patterns of sporophytic and gametophytic gene expression in the fern <i>Polypodium amorphum</i>, a member of the homosporous polypod lineage that comprises 80% of extant fern diversity, to assess how expression of a common genome is partitioned between two morphologically, ecologically, and nutritionally independent phases. Using RNA-sequencing, we generated transcriptome profiles for three replicates of paired samples of sporophyte leaf tissue and whole gametophytes to identify genes with significant differences in expression between the two phases. We found a nearly 90% overlap in the identity and expression levels of the genes expressed in both sporophytes and gametophytes, with less than 3% of genes uniquely expressed in either phase. We compare our results to those from similar studies to establish how phase-specific gene expression varies among major land plant lineages. Notably, despite having greater similarity in the identity of gene families shared between <i>P. amorphum</i> and angiosperms, <i>P. amorphum</i> has phase-specific gene expression profiles that are more like bryophytes and lycophytes than seed plants. Our findings suggest that shared patterns of phase-specific gene expression among seed-free plants likely reflect having relatively large, photosynthetic gametophytes (compared to the gametophytes of seed plants that are highly reduced). Phylogenetic analyses were used to further investigate the evolution of phase-specific expression for the phototropin, terpene synthase, and MADS-box gene families.}, Doi = {10.3389/fpls.2018.01450}, Key = {fds339280} } @article{fds322311, Author = {Rothfels, CJ and Pryer, KM and Li, F}, Title = {Next‐generation polyploid phylogenetics: Rapid resolution of hybrid polyploid complexes using PacBio single‐molecule sequencing}, Journal = {New Phytologist}, Volume = {213}, Number = {1}, Pages = {413-429}, Year = {2017}, url = {http://dx.doi.org/10.1111/nph.14111}, Abstract = {Difficulties in generating nuclear data for polyploids have impeded phylogenetic study of these groups. We describe a high-throughput protocol and an associated bioinformatics pipeline (Pipeline for Untangling Reticulate Complexes (Purc)) that is able to generate these data quickly and conveniently, and demonstrate its efficacy on accessions from the fern family Cystopteridaceae. We conclude with a demonstration of the downstream utility of these data by inferring a multi-labeled species tree for a subset of our accessions. We amplified four c. 1-kb-long nuclear loci and sequenced them in a parallel-tagged amplicon sequencing approach using the PacBio platform. Purc infers the final sequences from the raw reads via an iterative approach that corrects PCR and sequencing errors and removes PCR-mediated recombinant sequences (chimeras). We generated data for all gene copies (homeologs, paralogs, and segregating alleles) present in each of three sets of 50 mostly polyploid accessions, for four loci, in three PacBio runs (one run per set). From the raw sequencing reads, Purc was able to accurately infer the underlying sequences. This approach makes it easy and economical to study the phylogenetics of polyploids, and, in conjunction with recent analytical advances, facilitates investigation of broad patterns of polyploid evolution.}, Doi = {10.1111/nph.14111}, Key = {fds322311} } @article{fds323623, Author = {Hirai, RY and Schuettpelz, E and Huiet, L and Pryer, KM and Smith, AR and Prado, J}, Title = {Phylogeny and relationships of the neotropical Adiantum raddianum group (Pteridaceae)}, Journal = {Taxon}, Volume = {65}, Number = {6}, Pages = {1225-1235}, Publisher = {WILEY}, Year = {2016}, url = {http://dx.doi.org/10.12705/656.1}, Abstract = {With more than 200 species, the maidenhair fern genus Adiantum is among the top ten most diverse fern genera. Adiantum is pantropical in distribution and, due to the presence of a unique synapomorphy (sporangia borne on indusia rather than laminae), perhaps the most easily recognized fern genus. Many of its members, including numerous cultivars derived from A. raddianum, are grown as ornamentals. Because of its size, a comprehensive taxonomic study of Adiantum is difficult and the genus is perhaps better approached through a series of narrower studies. Here, we focus specifically on A. raddianum and putative allies. We find a newly defined A. raddianum group to be strongly supported as monophyletic and segregated from other maidenhair ferns on the basis of genetic as well as morphological characteristics. Bayesian inference and maximum likelihood analyses of plastid atpA, chlL, chlN, rbcL, and rpoA sequences support the A raddianum clade as sister to A poiretii and its allies. We identify round-reniform indusia to be a characteristic of the A.raddianum group (vs. lunate in the A.poiretii group). Additionally, we find species in the Apoiretii group to differ in having a unique 66 nucleotide deletion in our chlN gene alignment. The neotropical Araddianum group comprises at least 17 species (14 studied here), some widely distributed; one was recently described (A. alan-smithii).}, Doi = {10.12705/656.1}, Key = {fds323623} } @article{fds324241, Author = {Haufler, CH and Pryer, KM and Schuettpelz, E and Sessa, EB and Farrar, DR and Moran, R and Schneller, JJ and Watkins Jr and JE and Windham, MD}, Title = {Sex and the single gametophyte: Revising the homosporous vascular plant life cycle in light of contemporary research}, Journal = {BioScience}, Volume = {66}, Number = {11}, Pages = {928-937}, Publisher = {Oxford University Press}, Year = {2016}, url = {http://dx.doi.org/10.1093/biosci/biw108}, Abstract = {Homosporous vascular plants are typically depicted as extreme inbreeders, with bisexual gametophytes that produce strictly homozygous sporophytes. This view is promulgated in textbook life cycles despite ample evidence that natural populations of most species regularly outcross. We review research on a variety of mechanisms, including genetic load, asynchronous production of eggs and sperm, and pheromonal control of gamete production, that actively promote heterozygosity in ferns and lycophytes. Evolution of the land plants cannot be reconstructed without accurate depictions of the unique life cycle that has helped make ferns the second most diverse lineage of vascular plants on Earth. With revised illustrations and definitions, we provide scientists, educators, and students with a contemporary understanding of fern and lycophyte reproduction, revealing them as evolutionarily dynamic and exploiting a wide range of mating systems.}, Doi = {10.1093/biosci/biw108}, Key = {fds324241} } @article{fds324242, Author = {Li, F and Kuo, L and Pryer, KM and Rothfels, CJ}, Title = {Genes translocated into the plastid inverted repeat show decelerated substitution rates and elevated GC content}, Journal = {Genome Biology and Evolution}, Volume = {8}, Number = {8}, Pages = {2452-2458}, Publisher = {Oxford University Press}, Year = {2016}, url = {http://dx.doi.org/10.1093/gbe/evw167}, Abstract = {Plant chloroplast genomes (plastomes) are characterized by an inverted repeat (IR) region and two larger single copy (SC) regions. Patterns of molecular evolution in the IR and SC regions differ, most notably by a reduced rate of nucleotide substitution in the IR compared to the SC region. In addition, the organization and structure of plastomes is fluid, and rearrangements through time have repeatedly shuffled genes into and out of the IR, providing recurrent natural experiments on how chloroplast genome structure can impact rates and patterns of molecular evolution. Here we examine four loci (psbA, ycf2, rps7, and rps12 exon 2-3) that were translocated from the SC into the IR during fern evolution. We use a model-based method, within a phylogenetic context, to test for substitution rate shifts. All four loci show a significant, 2- to 3-fold deceleration in their substitution rate following translocation into the IR, a phenomenon not observed in any other, nontranslocated plastid genes. Also, we show that after translocation, the GC content of the third codon position and of the noncoding regions is significantly increased, implying that gene conversion within the IR is GC-biased. Taken together, our results suggest that the IR region not only reduces substitution rates, but also impacts nucleotide composition. This finding highlights a potential vulnerability of correlating substitution rate heterogeneity with organismal life history traits without knowledge of the underlying genome structure.}, Doi = {10.1093/gbe/evw167}, Key = {fds324242} } @article{fds322312, Author = {Schuettpelz, E and Chen, C and Kessler, M and Pinson, JB and Johnson, G and Davila, A and Cochran, AT and Huiet, L and Pryer, KM}, Title = {A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data}, Journal = {Taxon}, Volume = {65}, Number = {4}, Pages = {708-722}, Publisher = {WILEY}, Year = {2016}, url = {http://dx.doi.org/10.12705/654.2}, Abstract = {Vittarioid ferns compose a well-supported clade of 100-130 species of highly simplified epiphytes in the family Pteridaceae. Generic circumscriptions within the vittarioid clade were among the first in ferns to be evaluated and revised based on molecular phylogenetic data. Initial analyses of rbcL sequences revealed strong geographic structure and demonstrated that the two largest vittarioid genera, as then defined, each had phylogenetically distinct American and Old World components. The results of subsequent studies that included as many as 36 individuals of 33 species, but still relied on a single gene, were generally consistent with the early findings. Here, we build upon the previous datasets, incorporating many more samples (138 individuals representing 72 species) and additional plastid markers (atpA, chlN, rbcL, rpoA). Analysis of our larger dataset serves to better characterize known lineages, reveals new lineages, and ultimately uncovers an underlying geographic signal that is even stronger than was previously appreciated. In our revised generic classification, we recognize a total of eleven vittarioid genera. Each genus, including the new genus Antrophyopsis (Benedict) Schuettp., stat. nov., is readily diagnosable based on morphology, with micromorphological characters related to soral paraphyses and spores complementing more obvious features such as venation and the distribution of sporangia. A key to the currently recognized vittarioid genera, brief generic descriptions, and five new species combinations are provided.}, Doi = {10.12705/654.2}, Key = {fds322312} } @article{fds322313, Author = {Pryer, KM and Huiet, L and Li, F and Rothfels, CJ and Schuettpelz, E}, Title = {Maidenhair ferns, Adiantum, are indeed monophyletic and sister to shoestring ferns, vittarioids (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {41}, Number = {1}, Pages = {17-23}, Publisher = {American Society of Plant Taxonomists}, Year = {2016}, url = {http://dx.doi.org/10.1600/036364416x690660}, Abstract = {Across the tree of life, molecular phylogenetic studies often reveal surprising relationships between taxa with radically different morphologies that have long obscured their close affiliations. A spectacular botanical example is Rafflesia, a holoparasite that produces the largest flowers in the world, but that evolved from tiny-flowered ancestors within the Euphorbiaceae. Outside of parasitic lineages, such abrupt transformations are rarely seen. One exception involves the “maidenhair ferns” (Adiantum), which are quintessential ferns: beautifully dissected, terrestrial, and shade loving. The closely related “shoestring ferns” (vittarioids), in contrast, have an extremely simplified morphology, are canopy-dwelling epiphytes, and exhibit greatly accelerated rates of molecular evolution. While Adiantum and the vittarioids together have been shown to form a robust monophyletic group (adiantoids), there remain unanswered questions regarding the monophyly of Adiantum and the evolutionary history of the vittarioids. Here we review recent phylogenetic evidence suggesting support for the monophyly of Adiantum, and analyze new plastid data to confirm this result. We find that Adiantum is monophyletic and sister to the vittarioids. With this robust phylogenetic framework established for the broadest relationships in the adiantoid clade, we can now focus on understanding the evolutionary processes associated with the extreme morphological, ecological, and genetic transitions that took place within this lineage.}, Doi = {10.1600/036364416x690660}, Key = {fds322313} } @article{fds358011, Author = {Wolf, PG and Sessa, EB and Marchant, DB and Li, F and Rothfels, CJ and Sigel, EM and Gitzendanner, MA and Visger, CJ and Banks, JA and Soltis, DE}, Title = {An exploration into fern genome space}, Journal = {Genome Biology and Evolution}, Volume = {7}, Number = {9}, Pages = {2533-2544}, Publisher = {Oxford University Press}, Year = {2015}, Key = {fds358011} } @article{fds230052, Author = {Schuettpelz, E and Pryer, KM and Windham, MD}, Title = {A unified approach to taxonomic delimitation in the fern genus Pentagramma (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {40}, Number = {3}, Pages = {629-644}, Publisher = {American Society of Plant Taxonomists}, Year = {2015}, ISSN = {0363-6445}, url = {http://hdl.handle.net/10161/10796 Duke open access}, Abstract = {The native goldback and silverback ferns of western North America, composing the genus Pentagramma, are phylogenetically isolated within the xeric-adapted cheilanthoid clade. Although species-poor compared to its sister group, Pentagramma encompasses a diverse array of morphotypes, cytotypes, and flavonoid chemotypes. Because the differences are generally cryptic, however, the various entities are usually recognized at an infraspecific level. In recent years, as many as five subspecies have been ascribed to P. triangularis, and only P. pallida has been considered sufficiently divergent to warrant recognition as a distinct species. In this study, we take a unified approach to taxonomic delimitation in Pentagramma. Combining spore studies with phylogenetic analyses of plastid and nuclear sequences, we identify six genetically and morphologically distinct diploid lineages, each of which is here treated as a species. A new species is described (P. glanduloviscida) and three new combinations are made (P. maxonii, P. rebmanii, and P. viscosa).}, Doi = {10.1600/036364415x689366}, Key = {fds230052} } @article{fds230053, Author = {Li, F and Rothfels, CJ and Melkonian, M and Villarreal, JC and Stevenson, DW and Graham, SW and Wong, GK-S and Mathews, S and Pryer, KM}, Title = {The origin and evolution of phototropins}, Journal = {Frontiers in plant science}, Volume = {6}, Pages = {637}, Publisher = {Frontiers}, Year = {2015}, url = {http://hdl.handle.net/10161/10800 Duke open access}, Abstract = {Plant phototropism, the ability to bend toward or away from light, is predominantly controlled by blue-light photoreceptors, the phototropins. Although phototropins have been well-characterized in Arabidopsis thaliana, their evolutionary history is largely unknown. In this study, we complete an in-depth survey of phototropin homologs across land plants and algae using newly available transcriptomic and genomic data. We show that phototropins originated in an ancestor of Viridiplantae (land plants + green algae). Phototropins repeatedly underwent independent duplications in most major land-plant lineages (mosses, lycophytes, ferns, and seed plants), but remained single-copy genes in liverworts and hornworts-an evolutionary pattern shared with another family of photoreceptors, the phytochromes. Following each major duplication event, the phototropins differentiated in parallel, resulting in two specialized, yet partially overlapping, functional forms that primarily mediate either low- or high-light responses. Our detailed phylogeny enables us to not only uncover new phototropin lineages, but also link our understanding of phototropin function in Arabidopsis with what is known in Adiantum and Physcomitrella (the major model organisms outside of flowering plants). We propose that the convergent functional divergences of phototropin paralogs likely contributed to the success of plants through time in adapting to habitats with diverse and heterogeneous light conditions.}, Doi = {10.3389/fpls.2015.00637}, Key = {fds230053} } @article{fds230055, Author = {Huiet, L and Lenz, M and Nelson, JK and Pryer, KM and Smith, AR}, Title = {Adiantum shastense, a new species of maidenhair fern from California}, Journal = {PhytoKeys}, Number = {53}, Pages = {73}, Publisher = {Pensoft Publishers}, Year = {2015}, ISSN = {1314-2011}, url = {http://hdl.handle.net/10161/10799 Duke open access}, Abstract = {A new species of Adiantum is described from California. This species is endemic to northern California and is currently known only from Shasta County. We describe its discovery after first being collected over a century ago and distinguish it from Adiantumjordanii and Adiantumcapillus-veneris. It is evergreen and is sometimes, but not always, associated with limestone. The range of Adiantumshastense Huiet & A.R.Sm., sp. nov., is similar to several other Shasta County endemics that occur in the mesic forests of the Eastern Klamath Range, close to Shasta Lake, on limestone and metasedimentary substrates.}, Doi = {10.3897/phytokeys.53.5151}, Key = {fds230055} } @article{fds230056, Author = {Rothfels, CJ and Li, F and Sigel, EM and Huiet, L and Larsson, A and Burge, DO and Ruhsam, M and Deyholos, M and Soltis, DE and Stewart Jr, CN}, Title = {The evolutionary history of ferns inferred from 25 low‐copy nuclear genes}, Journal = {American Journal of Botany}, Volume = {102}, Number = {7}, Pages = {1089-1107}, Publisher = {Botanical Society of America}, Year = {2015}, ISSN = {0002-9122}, url = {http://hdl.handle.net/10161/10798 Duke open access}, Abstract = {<h4>Unlabelled</h4>•<h4>Premise of the study</h4>Understanding fern (monilophyte) phylogeny and its evolutionary timescale is critical for broad investigations of the evolution of land plants, and for providing the point of comparison necessary for studying the evolution of the fern sister group, seed plants. Molecular phylogenetic investigations have revolutionized our understanding of fern phylogeny, however, to date, these studies have relied almost exclusively on plastid data.•<h4>Methods</h4>Here we take a curated phylogenomics approach to infer the first broad fern phylogeny from multiple nuclear loci, by combining broad taxon sampling (73 ferns and 12 outgroup species) with focused character sampling (25 loci comprising 35877 bp), along with rigorous alignment, orthology inference and model selection.•<h4>Key results</h4>Our phylogeny corroborates some earlier inferences and provides novel insights; in particular, we find strong support for Equisetales as sister to the rest of ferns, Marattiales as sister to leptosporangiate ferns, and Dennstaedtiaceae as sister to the eupolypods. Our divergence-time analyses reveal that divergences among the extant fern orders all occurred prior to ∼200 MYA. Finally, our species-tree inferences are congruent with analyses of concatenated data, but generally with lower support. Those cases where species-tree support values are higher than expected involve relationships that have been supported by smaller plastid datasets, suggesting that deep coalescence may be reducing support from the concatenated nuclear data.•<h4>Conclusions</h4>Our study demonstrates the utility of a curated phylogenomics approach to inferring fern phylogeny, and highlights the need to consider underlying data characteristics, along with data quantity, in phylogenetic studies.}, Doi = {10.3732/ajb.1500089}, Key = {fds230056} } @article{fds230057, Author = {Melkonian, M and Pryer, KM and Rothfels, CJ and Graham, SW and Li, F and Villarreal, JC and Wong, GK and Stevenson, DW and Mathews, S}, Title = {Phytochrome diversity in green plants and the origin of canonical plant phytochromes}, Volume = {6}, Number = {1}, Pages = {1-12}, Publisher = {Nature Publishing Group}, Year = {2015}, url = {http://hdl.handle.net/10161/10330 Duke open access}, Abstract = {Phytochromes are red/far-red photoreceptors that play essential roles in diverse plant morphogenetic and physiological responses to light. Despite their functional significance, phytochrome diversity and evolution across photosynthetic eukaryotes remain poorly understood. Using newly available transcriptomic and genomic data we show that canonical plant phytochromes originated in a common ancestor of streptophytes (charophyte algae and land plants). Phytochromes in charophyte algae are structurally diverse, including canonical and non-canonical forms, whereas in land plants, phytochrome structure is highly conserved. Liverworts, hornworts and Selaginella apparently possess a single phytochrome, whereas independent gene duplications occurred within mosses, lycopods, ferns and seed plants, leading to diverse phytochrome families in these clades. Surprisingly, the phytochrome portions of algal and land plant neochromes, a chimera of phytochrome and phototropin, appear to share a common origin. Our results reveal novel phytochrome clades and establish the basis for understanding phytochrome functional evolution in land plants and their algal relatives.}, Doi = {10.1038/ncomms8852}, Key = {fds230057} } @article{fds230060, Author = {Rothfels, CJ and Johnson, AK and Hovenkamp, PH and Swofford, DL and Roskam, HC and Fraser-Jenkins, CR and Windham, MD and Pryer, KM}, Title = {Natural hybridization between genera that diverged from each other approximately 60 million years ago}, Journal = {The American Naturalist}, Volume = {185}, Number = {3}, Pages = {433-442}, Publisher = {University of Chicago Press Chicago, IL}, Year = {2015}, ISSN = {0003-0147}, url = {http://hdl.handle.net/10161/10248 Duke open access}, Abstract = {A fern from the French Pyrenees-×Cystocarpium roskamianum-is a recently formed intergeneric hybrid between parental lineages that diverged from each other approximately 60 million years ago (mya; 95% highest posterior density: 40.2-76.2 mya). This is an extraordinarily deep hybridization event, roughly akin to an elephant hybridizing with a manatee or a human with a lemur. In the context of other reported deep hybrids, this finding suggests that populations of ferns, and other plants with abiotically mediated fertilization, may evolve reproductive incompatibilities more slowly, perhaps because they lack many of the premating isolation mechanisms that characterize most other groups of organisms. This conclusion implies that major features of Earth's biodiversity-such as the relatively small number of species of ferns compared to those of angiosperms-may be, in part, an indirect by-product of this slower "speciation clock" rather than a direct consequence of adaptive innovations by the more diverse lineages.}, Doi = {10.1086/679662}, Key = {fds230060} } @article{fds376514, Author = {Li, F and Melkonian, M and Rothfels, CJ and Villarreal, JC and Stevenson, DW and Graham, SW and Wong, GK and Pryer, KM and Mathews, S}, Title = {Phytochrome diversity in green plants and the origin of canonical plant phytochromes}, Journal = {Nature communications}, Volume = {6}, Number = {1}, Pages = {1-12}, Publisher = {Nature Publishing Group}, Year = {2015}, Key = {fds376514} } @article{fds322315, Author = {Kao, T and Pryer, KM and Turner, MD and White, RA and Korall, P}, Title = {Origins of the endemic scaly tree ferns on the Galápagos and Cocos Islands}, Journal = {International Journal of Plant Sciences}, Volume = {176}, Number = {9}, Pages = {869-879}, Publisher = {University of Chicago Press Chicago, IL}, Year = {2015}, url = {http://dx.doi.org/10.1086/683303}, Abstract = {Premise of research. Successful long-distance dispersal is rarely observed in scaly tree ferns (Cyatheaceae). Nevertheless, recent molecular evidence has suggested that the four endemic scaly tree ferns on the Galápagos Archipelago (Cyathea weatherbyana) and Cocos Island (Cyathea alfonsiana, Cyathea nesiotica, and Cyathea notabilis), two oceanic island groups west of Central and northern South America, probably each originated from different mainland America ancestors. However, the phylogenetic relationships inferred among these endemics and their mainland relatives have been unclear. This study is aimed at better resolving the relationships and tracing the origins of these island endemics. Methodology. Five plastid regions from 35 Cyathea species were analyzed to reconstruct phylogenetic relationships using parsimony, likelihood, and Bayesian approaches. We also estimated divergence times of these species, and our chronogram was used to reconstruct their biogeographical range history. Pivotal results. Our well-resolved phylogenetic tree of Cyathea, which is in agreement with previous studies, shows that when the four Galápagos and Cocos endemics are included, they each belong to separate subclades. Our biogeographical study suggests that the four endemics originated from independent colonization events from mainland America and that there was no dispersal of Cyathea between the island groups. We reveal more detailed relationships among the endemics and their respective close mainland relatives; some of these relationships differ from previous studies. Our findings are corroborated by new morphological data from ongoing stem anatomy studies. Conclusions. The four scaly tree ferns endemic to the Galápagos and Cocos Islands each did indeed originate as independent colonization events from separate sources in mainland America, and their closest relatives are identified here.}, Doi = {10.1086/683303}, Key = {fds322315} } @article{fds322314, Author = {Grusz, AL and Pryer, KM}, Title = {Development of microsatellite markers for the apomictic triploid fern Myriopteris lindheimeri (Pteridaceae)}, Journal = {Applications in plant sciences}, Volume = {3}, Number = {11}, Pages = {1500061}, Publisher = {Botanical Society of America}, Year = {2015}, url = {http://dx.doi.org/10.3732/apps.1500061}, Abstract = {<h4>Premise of the study</h4>Microsatellite markers were developed for investigating the population dynamics of Myriopteris lindheimeri (Pteridaceae), an apomictic triploid fern endemic to deserts of the southwestern United States and Mexico.<h4>Methods and results</h4>Using 454 sequencing, 21 microsatellite markers were developed. Of these, 14 were polymorphic with up to five alleles per locus and eight markers amplified in one or more congeneric close relatives (M. covillei, M. fendleri, M. aurea, and M. rufa). To demonstrate marker utility, M. lindheimeri samples from three Arizona populations were genotyped at nine loci. For each population, diversity measures including percent polymorphic loci, frequency of heterozygotes across all loci, and genotypic diversity were calculated. Across the three populations, on average, 63% of loci were polymorphic, the average frequency of heterozygotes (across all loci) was 0.32, and average genotypic diversity was 0.34.<h4>Conclusions</h4>These markers provide a foundation for future studies exploring polyploidy and apomixis in myriopterid ferns.}, Doi = {10.3732/apps.1500061}, Key = {fds322314} } @article{fds353345, Author = {Rothfels, CJ and Wong, GK and Stevenson, DW and Li, F and Mathews, S and Melkonian, M and Pryer, KM and Graham, SW and Villarreal, JC}, Title = {The Origin and Evolution of Phototropins}, Year = {2015}, Key = {fds353345} } @article{fds230063, Author = {Sessa, EB and Banks, JA and Barker, MS and Der, JP and Duffy, AM and Graham, SW and Hasebe, M and Langdale, J and Li, F-W and Marchant, DB and Pryer, KM and Rothfels, CJ and Roux, SJ and Salmi, ML and Sigel, EM and Soltis, DE and Soltis, PS and Stevenson, DW and Wolf, PG}, Title = {Between two fern genomes}, Journal = {Gigascience}, Volume = {3}, Number = {1}, Pages = {15-15}, Publisher = {Oxford University Press}, Year = {2014}, Month = {September}, url = {http://hdl.handle.net/10161/10249 Duke open access}, Abstract = {Ferns are the only major lineage of vascular plants not represented by a sequenced nuclear genome. This lack of genome sequence information significantly impedes our ability to understand and reconstruct genome evolution not only in ferns, but across all land plants. Azolla and Ceratopteris are ideal and complementary candidates to be the first ferns to have their nuclear genomes sequenced. They differ dramatically in genome size, life history, and habit, and thus represent the immense diversity of extant ferns. Together, this pair of genomes will facilitate myriad large-scale comparative analyses across ferns and all land plants. Here we review the unique biological characteristics of ferns and describe a number of outstanding questions in plant biology that will benefit from the addition of ferns to the set of taxa with sequenced nuclear genomes. We explain why the fern clade is pivotal for understanding genome evolution across land plants, and we provide a rationale for how knowledge of fern genomes will enable progress in research beyond the ferns themselves.}, Doi = {10.1186/2047-217x-3-15}, Key = {fds230063} } @article{fds230064, Author = {Li, F-W and Pryer, KM}, Title = {Crowdfunding the Azolla fern genome project: a grassroots approach.}, Journal = {GigaScience}, Volume = {3}, Number = {1}, Pages = {16}, Publisher = {Oxford University Press}, Year = {2014}, Month = {January}, url = {http://hdl.handle.net/10161/10250 Duke open access}, Abstract = {Much of science progresses within the tight boundaries of what is often seen as a "black box". Though familiar to funding agencies, researchers and the academic journals they publish in, it is an entity that outsiders rarely get to peek into. Crowdfunding is a novel means that allows the public to participate in, as well as to support and witness advancements in science. Here we describe our recent crowdfunding efforts to sequence the Azolla genome, a little fern with massive green potential. Crowdfunding is a worthy platform not only for obtaining seed money for exploratory research, but also for engaging directly with the general public as a rewarding form of outreach.}, Doi = {10.1186/2047-217x-3-16}, Key = {fds230064} } @article{fds230054, Author = {Wolf, PG and Sessa, EB and Marchant, DB and Li, F and Rothfels, CJ and Sigel, EM and Gitzendanner, MA and Visger, CJ and Banks, JA and Soltis, DE}, Title = {An Exploration of Fern Genome Space}, Volume = {7}, Number = {9}, Pages = {2533-2544}, Publisher = {Botanical Society of America}, Year = {2014}, url = {http://hdl.handle.net/10161/10797 Duke open access}, Abstract = {Ferns are one of the few remaining major clades of land plants for which a complete genome sequence is lacking. Knowledge of genome space in ferns will enable broad-scale comparative analyses of land plant genes and genomes, provide insights into genome evolution across green plants, and shed light on genetic and genomic features that characterize ferns, such as their high chromosome numbers and large genome sizes. As part of an initial exploration into fern genome space, we used a whole genome shotgun sequencing approach to obtain low-density coverage (∼0.4X to 2X) for six fern species from the Polypodiales (Ceratopteris, Pteridium, Polypodium, Cystopteris), Cyatheales (Plagiogyria), and Gleicheniales (Dipteris). We explore these data to characterize the proportion of the nuclear genome represented by repetitive sequences (including DNA transposons, retrotransposons, ribosomal DNA, and simple repeats) and protein-coding genes, and to extract chloroplast and mitochondrial genome sequences. Such initial sweeps of fern genomes can provide information useful for selecting a promising candidate fern species for whole genome sequencing. We also describe variation of genomic traits across our sample and highlight some differences and similarities in repeat structure between ferns and seed plants.}, Doi = {10.1093/gbe/evv163}, Key = {fds230054} } @article{fds230061, Author = {Rothfels, CJ and Johnson, AK and Windham, MD and Pryer, KM}, Title = {Low‐copy nuclear data confirm rampant allopolyploidy in the Cystopteridaceae (Polypodiales)}, Journal = {Taxon}, Volume = {63}, Number = {5}, Pages = {1026-1036}, Publisher = {International Association for Plant Taxonomy}, Year = {2014}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.12705/635.32}, Abstract = {Here we present the first nuclear phylogeny for Cystopteridaceae (Polypodiales), using the single-copy locus gapCp “short”. This phylogeny corroborates broad results from plastid data in demonstrating strong support for the monophyly of the family’s three genera—Cystopteris, Acystopteris, and Gymnocarpium—and of the major groups within Cystopteris (C. montana, the sudetica and bulbifera clades, and the C. fragilis complex). In addition, it confirms the rampant hybridization (allopolyploidy) that has long been suspected within both Cystopteris and Gymnocarpium. In some cases, these data provide the first DNA-sequence-based evidence for previous hypotheses of polyploid species origins (such as the cosmopolitan G. dryopteris being an allotetraploid derivative of the diploids G. appalachianum and G. disjunctum). Most of the allopolyploids, however, have no formal taxonomic names. This pattern is particularly strong within the C. fragilis complex, where our results imply that the eight included accessions of “C. fragilis” represent at least six distinct allopolyploid taxa.}, Doi = {10.12705/635.32}, Key = {fds230061} } @article{fds230062, Author = {Sigel, EM and Windham, MD and Haufler, CH and Pryer, KM}, Title = {Phylogeny, divergence time estimates, and phylogeography of the diploid species of the Polypodium vulgare complex (Polypodiaceae)}, Journal = {Systematic Botany}, Volume = {39}, Number = {4}, Pages = {1042-1055}, Publisher = {American Society of Plant Taxonomists}, Year = {2014}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364414x683921}, Abstract = {The Polypodium vulgare complex (Polypodiaceae) comprises a well-studied group of fern taxa whose members are cryptically differentiated morphologically and have generated a confusing and highly reticulate species cluster. Once considered a single species spanning much of northern Eurasia and North America, P. vulgare has been segregated into 17 diploid and polyploid taxa as a result of cytotaxonomic work, hybridization experiments, and isozyme studies conducted during the 20th century. Despite considerable effort, however, the evolutionary relationships among the diploid members of the P. vulgare complex remain poorly resolved. Here we infer a diploids-only phylogeny of the P. vulgare complex and related species to test previous hypotheses concerning relationships within Polypodium sensu stricto. Using sequence data from four plastid loci (atpA, rbcL, matK, and trnG-trnR), we recovered a monophyletic P. vulgare complex comprising four well-supported clades. The P. vulgare complex is resolved as sister to the Neotropical P. plesiosorum group and these, in turn, are sister to the Asian endemic Pleurosoriopsis makinoi. Using divergence time analyses incorporating previously derived age constraints and fossil data, we estimate an early Miocene origin for the P. vulgare complex and a late Miocene-Pliocene origin for the four major diploid lineages of the complex, with the majority of extant diploid species diversifying from the late Miocene through the Pleistocene. Finally, we use our node age estimates to reassess previous hypotheses, and to propose new hypotheses, about the historical events that shaped the diversity and current geographic distribution of the diploid species of the P. vulgare complex.}, Doi = {10.1600/036364414x683921}, Key = {fds230062} } @article{fds230065, Author = {Sigel, EM and Windham, MD and Pryer, KM}, Title = {Evidence for reciprocal origins in Polypodium hesperium (Polypodiaceae): A fern model system for investigating how multiple origins shape allopolyploid genomes}, Journal = {American Journal of Botany}, Volume = {101}, Number = {9}, Pages = {1476-1485}, Publisher = {Botanical Society of America}, Year = {2014}, ISSN = {0002-9122}, url = {http://dx.doi.org/10.3732/ajb.1400190}, Abstract = {<h4>Unlabelled</h4>•<h4>Premise of the study</h4>Many polyploid species are composed of distinct lineages originating from multiple, independent polyploidization events. In the case of allopolyploids, reciprocal crosses between the same progenitor species can yield lineages with different uniparentally inherited plastid genomes. While likely common, there are few well-documented examples of such reciprocal origins. Here we examine a case of reciprocal allopolyploid origins in the fern Polypodium hesperium and present it as a natural model system for investigating the evolutionary potential of duplicated genomes.•<h4>Methods</h4>Using a combination of uniparentally inherited plastid and biparentally inherited nuclear sequence data, we investigated the distributions and relative ages of reciprocally formed lineages in Polypodium hesperium, an allotetraploid fern that is broadly distributed in western North America.•<h4>Key results</h4>The reciprocally derived plastid haplotypes of Polypodium hesperium are allopatric, with populations north and south of 42°N latitude having different plastid genomes. Incorporating biogeographic information and previously estimated ages for the diversification of its diploid progenitors, we estimate middle to late Pleistocene origins of P. hesperium.•<h4>Conclusions</h4>Several features of Polypodium hesperium make it a particularly promising system for investigating the evolutionary consequences of allopolyploidy. These include reciprocally derived lineages with disjunct geographic distributions, recent time of origin, and extant diploid progenitors.}, Doi = {10.3732/ajb.1400190}, Key = {fds230065} } @article{fds230067, Author = {Grusz, AL and Windham, MD and Yatskievych, G and Huiet, L and Gastony, GJ and Pryer, KM}, Title = {Patterns of diversification in the xeric-adapted fern genus Myriopteris (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {39}, Number = {3}, Pages = {698-714}, Publisher = {American Society of Plant Taxonomists}, Year = {2014}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364414x681518}, Abstract = {Strong selective pressures imposed by drought-prone habitats have contributed to extensive morphological convergence among the 400 + species of cheilanthoid ferns (Pteridaceae). As a result, generic circumscriptions based exclusively on macromorphology often prove to be non-monophyletic. Ongoing molecular phylogenetic analyses are providing the foundation for a revised classification of this challenging group and have begun to clarify its complex evolutionary history. As part of this effort, we generated and analyzed DNA sequence data for three plastid loci (rbcL, atpA, and the intergenic spacer trnG-trnR) for the myriopterid clade, one of the largest monophyletic groups of cheilanthoid ferns. This lineage encompasses 47 primarily North and Central American taxa previously included in Cheilanthes but now placed in the recircumscribed genus Myriopteris. Here, we infer a phylogeny for the group and examine key morphological characters across this phylogeny. We also include a brief discussion of the three well-supported Myriopteris subclades, along with a review of reproductive mode and known ploidy levels for members of this early diverging lineage of cheilanthoid ferns.}, Doi = {10.1600/036364414x681518}, Key = {fds230067} } @article{fds230068, Author = {Li, F and Villarreal, JC and Kelly, S and Rothfels, CJ and Melkonian, M and Frangedakis, E and Ruhsam, M and Sigel, EM and Der, JP and Pittermann, J}, Title = {Horizontal transfer of an adaptive chimeric photoreceptor from bryophytes to ferns}, Journal = {Proceedings of the National Academy of Sciences}, Volume = {111}, Number = {18}, Pages = {6672-6677}, Publisher = {National Academy of Sciences}, Year = {2014}, ISSN = {0027-8424}, url = {http://dx.doi.org/10.1073/pnas.1319929111}, Abstract = {Ferns are well known for their shade-dwelling habits. Their ability to thrive under low-light conditions has been linked to the evolution of a novel chimeric photoreceptor--neochrome--that fuses red-sensing phytochrome and blue-sensing phototropin modules into a single gene, thereby optimizing phototropic responses. Despite being implicated in facilitating the diversification of modern ferns, the origin of neochrome has remained a mystery. We present evidence for neochrome in hornworts (a bryophyte lineage) and demonstrate that ferns acquired neochrome from hornworts via horizontal gene transfer (HGT). Fern neochromes are nested within hornwort neochromes in our large-scale phylogenetic reconstructions of phototropin and phytochrome gene families. Divergence date estimates further support the HGT hypothesis, with fern and hornwort neochromes diverging 179 Mya, long after the split between the two plant lineages (at least 400 Mya). By analyzing the draft genome of the hornwort Anthoceros punctatus, we also discovered a previously unidentified phototropin gene that likely represents the ancestral lineage of the neochrome phototropin module. Thus, a neochrome originating in hornworts was transferred horizontally to ferns, where it may have played a significant role in the diversification of modern ferns.}, Doi = {10.1073/pnas.1319929111}, Key = {fds230068} } @article{fds230069, Author = {Sigel, EM and Windham, MD and Smith, AR and Dyer, RJ and Pryer, KM}, Title = {Rediscovery of Polypodium calirhiza (Polypodiaceae) in Mexico}, Journal = {Brittonia}, Volume = {66}, Number = {3}, Pages = {278-286}, Publisher = {Springer US}, Year = {2014}, ISSN = {0007-196X}, url = {http://dx.doi.org/10.1007/s12228-014-9332-6}, Abstract = {This study addresses reported discrepancies regarding the occurrence of Polypodium calirhiza in Mexico. The original paper describing this taxon cited collections from Mexico, but the species was omitted from the recent Pteridophytes of Mexico. Originally treated as a tetraploid cytotype of P. californicum, P. calirhiza now is hypothesized to have arisen through hybridization between P. glycyrrhiza and P. californicum. The tetraploid can be difficult to distinguish from either of its putative parents, but especially so from P. californicum. Our analyses show that a combination of spore length and abaxial rachis scale morphology consistently distinguishes P. calirhiza from P. californicum, and we confirm that both species occur in Mexico. Although occasionally found growing together in the United States, the two species are strongly allopatric in Mexico: P. californicum is restricted to coastal regions of the Baja California peninsula and neighboring Pacific islands, whereas P. calirhiza grows at high elevations in central and southern Mexico. The occurrence of P. calirhiza in Oaxaca, Mexico, marks the southernmost extent of the P. vulgare complex in the Western Hemisphere. © 2014 The New York Botanical Garden.}, Doi = {10.1007/s12228-014-9332-6}, Key = {fds230069} } @article{fds304306, Author = {Korall, P and Pryer, KM}, Title = {Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal}, Journal = {Journal of biogeography}, Volume = {41}, Number = {2}, Pages = {402-413}, Year = {2014}, ISSN = {0305-0270}, url = {http://dx.doi.org/10.1111/jbi.12222}, Abstract = {Aim: Scaly tree ferns, Cyatheaceae, are a well-supported group of mostly tree-forming ferns found throughout the tropics, the subtropics and the south-temperate zone. Fossil evidence shows that the lineage originated in the Late Jurassic period. We reconstructed large-scale historical biogeographical patterns of Cyatheaceae and tested the hypothesis that some of the observed distribution patterns are in fact compatible, in time and space, with a vicariance scenario related to the break-up of Gondwana. Location: Tropics, subtropics and south-temperate areas of the world. Methods: The historical biogeography of Cyatheaceae was analysed in a maximum likelihood framework using Lagrange. The 78 ingroup taxa are representative of the geographical distribution of the entire family. The phylogenies that served as a basis for the analyses were obtained by Bayesian inference analyses of mainly previously published DNA sequence data using MrBayes. Lineage divergence dates were estimated in a Bayesian Markov chain Monte Carlo framework using beast. Results: Cyatheaceae originated in the Late Jurassic in either South America or Australasia. Following a range expansion, the ancestral distribution of the marginate-scaled clade included both these areas, whereas Sphaeropteris is reconstructed as having its origin only in Australasia. Within the marginate-scaled clade, reconstructions of early divergences are hampered by the unresolved relationships among the Alsophila, Cyathea and Gymnosphaera lineages. Nevertheless, it is clear that the occurrence of the Cyathea and Sphaeropteris lineages in South America may be related to vicariance, whereas transoceanic dispersal needs to be inferred for the range shifts seen in Alsophila and Gymnosphaera. Main conclusions: The evolutionary history of Cyatheaceae involves both Gondwanan vicariance scenarios as well as long-distance dispersal events. The number of transoceanic dispersals reconstructed for the family is rather few when compared with other fern lineages. We suggest that a causal relationship between reproductive mode (outcrossing) and dispersal limitations is the most plausible explanation for the pattern observed. © 2013 The Authors Journal of Biogeography Published by John Wiley & Sons Ltd.}, Doi = {10.1111/jbi.12222}, Key = {fds304306} } @article{fds353382, Author = {Li, F and Pryer, KM}, Title = {Crowdfunding the Azolla fern genome project: a grassroots approach}, Journal = {GigaScience}, Volume = {3}, Number = {1}, Pages = {2047-217X}, Publisher = {Oxford University Press}, Year = {2014}, Key = {fds353382} } @article{fds353383, Author = {Sessa, EB and Banks, JA and Barker, MS and Der, JP and Duffy, AM and Graham, SW and Hasebe, M and Langdale, J and Li, F and Marchant, DB}, Title = {Between two fern genomes}, Journal = {GigaScience}, Volume = {3}, Number = {1}, Pages = {2047-217X}, Publisher = {Oxford University Press}, Year = {2014}, Key = {fds353383} } @article{fds230071, Author = {Rothfels, CJ and Larsson, A and Li, F and Sigel, EM and Huiet, L and Burge, DO and Ruhsam, M and Graham, SW and Stevenson, DW and Wong, GK}, Title = {Transcriptome-mining for single-copy nuclear markers in ferns}, Journal = {PloS one}, Volume = {8}, Number = {10}, Pages = {e76957}, Publisher = {Public Library of Science}, Year = {2013}, url = {http://www.ncbi.nlm.nih.gov/pubmed/24116189}, Abstract = {<h4>Background</h4>Molecular phylogenetic investigations have revolutionized our understanding of the evolutionary history of ferns-the second-most species-rich major group of vascular plants, and the sister clade to seed plants. The general absence of genomic resources available for this important group of plants, however, has resulted in the strong dependence of these studies on plastid data; nuclear or mitochondrial data have been rarely used. In this study, we utilize transcriptome data to design primers for nuclear markers for use in studies of fern evolutionary biology, and demonstrate the utility of these markers across the largest order of ferns, the Polypodiales.<h4>Principal findings</h4>We present 20 novel single-copy nuclear regions, across 10 distinct protein-coding genes: ApPEFP_C, cryptochrome 2, cryptochrome 4, DET1, gapCpSh, IBR3, pgiC, SQD1, TPLATE, and transducin. These loci, individually and in combination, show strong resolving power across the Polypodiales phylogeny, and are readily amplified and sequenced from our genomic DNA test set (from 15 diploid Polypodiales species). For each region, we also present transcriptome alignments of the focal locus and related paralogs-curated broadly across ferns-that will allow researchers to develop their own primer sets for fern taxa outside of the Polypodiales. Analyses of sequence data generated from our genomic DNA test set reveal strong effects of partitioning schemes on support levels and, to a much lesser extent, on topology. A model partitioned by codon position is strongly favored, and analyses of the combined data yield a Polypodiales phylogeny that is well-supported and consistent with earlier studies of this group.<h4>Conclusions</h4>The 20 single-copy regions presented here more than triple the single-copy nuclear regions available for use in ferns. They provide a much-needed opportunity to assess plastid-derived hypotheses of relationships within the ferns, and increase our capacity to explore aspects of fern evolution previously unavailable to scientific investigation.}, Doi = {10.1371/journal.pone.0076957}, Key = {fds230071} } @article{fds230103, Author = {Rothfels, CJ and Windham, MD and Pryer, KM}, Title = {A plastid phylogeny of the cosmopolitan fern family Cystopteridaceae (Polypodiopsida)}, Journal = {Systematic Botany}, Volume = {38}, Number = {2}, Pages = {295-306}, Publisher = {American Society of Plant Taxonomists}, Year = {2013}, url = {http://dx.doi.org/10.1600/036364413x666787}, Abstract = {Among the novel results of recent molecular phylogenetic analyses are the unexpectedly close evolutionary relationships of the genera Acystopteris, Cystopteris, and Gymnocarpium, and the phylogenetic isolation of these genera from Woodsia. As a consequence, these three genera have been removed from Woodsiaceae and placed into their own family, the Cystopteridaceae. Despite the ubiquity of this family in rocky habitats across the northern hemisphere, and its cosmopolitan distribution (occurring on every continent except Antarctica), sampling of the Cystopteridaceae in phylogenetic studies to date has been sparse. Here we assemble a three-locus plastid dataset (matK, rbcL, trnG-R) that includes most recognized species in the family and multiple accessions of widespread taxa from across their geographic ranges. All three sampled genera are robustly supported as monophyletic, Cystopteris is strongly supported as sister to Acystopteris, and those two genera together are sister to Gymnocarpium. The Gymnocarpium phylogeny is deeply divided into three major clades, which we label the disjunctum clade, the robertianum clade, and core Gymnocarpium. The Cystopteris phylogeny, similarly, features four deeply diverged clades: C. montana, the sudetica clade, the bulbifera clade, and the fragilis complex. Acystopteris includes only three species, each of which is supported as monophyletic, with A. taiwaniana sister to the japonica/tenuisecta clade. Our results yield the first species-level phylogeny of the Cystopteridaceae and the first molecular phylogenetic evidence for species boundaries. These data provide an essential foundation for further investigations of complex patterns of geographic diversification, speciation, and reticulation in this family.}, Doi = {10.1600/036364413x666787}, Key = {fds230103} } @article{fds230105, Author = {León, B and Rothfels, CJ and Arakaki, M and Young, KR and Pryer, KM}, Title = {Revealing a cryptic fern distribution through DNA sequencing: Pityrogramma trifoliata in the Western Andes of Peru}, Journal = {American Fern Journal}, Volume = {103}, Number = {1}, Pages = {40-48}, Publisher = {The American Fern Society}, Year = {2013}, url = {http://dx.doi.org/10.1640/0002-8444-103.1.40}, Abstract = {Fern identification usually requires the use of mature sporophytes, since attempts to identify juveniles using morphological traits often provides unsatisfactory results. Here we examined young sporophytes found among boulders in a river basin of a xeric valley in central Peru. Attempts to identify these sporophytes first pointed to four different genera, two in Pteridaceae (Anogramma and Pityrogramma), and the others in Aspleniaceae (Asplenium) and Cystopteridaceae (Cystopteris). Here, we resolved this puzzle combining morphology and sequences of DNA (rbcL and trnG-R) that point to Pityrogramma trifoliata of Pteridaceae.}, Doi = {10.1640/0002-8444-103.1.40}, Key = {fds230105} } @article{fds230109, Author = {Grusz, AL and Pryer, KM and Yatskievych, G and Huiet, RL and Gastony, GJ and Windham, MD}, Title = {Refining the phylogeny of cheilanthoid ferns: The resurrection and recircumscription of Myriopteris (Pteridaceae)}, Journal = {Systematic Botany}, Year = {2013}, Key = {fds230109} } @article{fds230110, Author = {Korall, P and Pryer, KM}, Title = {Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal}, Journal = {Journal of Biogeography}, Volume = {40}, Number = {2}, Pages = {in press}, Year = {2013}, ISSN = {0305-0270}, url = {http://dx.doi.org/10.1111/jbi.12222}, Abstract = {<h4>Aim</h4>Scaly tree ferns, Cyatheaceae, are a well-supported group of mostly tree-forming ferns found throughout the tropics, the subtropics and the south-temperate zone. Fossil evidence shows that the lineage originated in the Late Jurassic period. We reconstructed large-scale historical biogeographical patterns of Cyatheaceae and tested the hypothesis that some of the observed distribution patterns are in fact compatible, in time and space, with a vicariance scenario related to the break-up of Gondwana.<h4>Location</h4>Tropics, subtropics and south-temperate areas of the world.<h4>Methods</h4>The historical biogeography of Cyatheaceae was analysed in a maximum likelihood framework using Lagrange. The 78 ingroup taxa are representative of the geographical distribution of the entire family. The phylogenies that served as a basis for the analyses were obtained by Bayesian inference analyses of mainly previously published DNA sequence data using MrBayes. Lineage divergence dates were estimated in a Bayesian Markov chain Monte Carlo framework using beast.<h4>Results</h4>Cyatheaceae originated in the Late Jurassic in either South America or Australasia. Following a range expansion, the ancestral distribution of the marginate-scaled clade included both these areas, whereas <i>Sphaeropteris</i> is reconstructed as having its origin only in Australasia. Within the marginate-scaled clade, reconstructions of early divergences are hampered by the unresolved relationships among the <i>Alsophila</i>, <i>Cyathea</i> and <i>Gymnosphaera</i> lineages. Nevertheless, it is clear that the occurrence of the <i>Cyathea</i> and <i>Sphaeropteris</i> lineages in South America may be related to vicariance, whereas transoceanic dispersal needs to be inferred for the range shifts seen in <i>Alsophila</i> and <i>Gymnosphaera</i>.<h4>Main conclusions</h4>The evolutionary history of Cyatheaceae involves both Gondwanan vicariance scenarios as well as long-distance dispersal events. The number of transoceanic dispersals reconstructed for the family is rather few when compared with other fern lineages. We suggest that a causal relationship between reproductive mode (outcrossing) and dispersal limitations is the most plausible explanation for the pattern observed.}, Doi = {10.1111/jbi.12222}, Key = {fds230110} } @article{fds230106, Author = {Beck, JB and Allison, JR and Pryer, KM and Windham, MD}, Title = {Identifying multiple origins of polyploid taxa: a multilocus study of the hybrid cloak fern (Astrolepis integerrima; Pteridaceae).}, Journal = {American journal of botany}, Volume = {99}, Number = {11}, Pages = {1857-1865}, Year = {2012}, Month = {November}, url = {http://www.ncbi.nlm.nih.gov/pubmed/23108464}, Abstract = {<h4>Premise of the study</h4>Molecular studies have shown that multiple origins of polyploid taxa are the rule rather than the exception. To understand the distribution and ecology of polyploid species and the evolutionary significance of polyploidy in general, it is important to delineate these independently derived lineages as accurately as possible. Although gene flow among polyploid lineages and backcrossing to their diploid parents often confound this process, such post origin gene flow is very infrequent in asexual polyploids. In this study, we estimate the number of independent origins of the apomictic allopolyploid fern Astrolepis integerrima, a morphologically heterogeneous species most common in the southwestern United States and Mexico, with outlying populations in the southeastern United States and the Caribbean.<h4>Methods</h4>Plastid DNA sequence and AFLP data were obtained from 33 A. integerrima individuals. Phylogenetic analysis of the sequence data and multidimensional clustering of the AFLP data were used to identify independently derived lineages.<h4>Key results</h4>Analysis of the two datasets identified 10 genetic groups within the 33 analyzed samples. These groups suggest a minimum of 10 origins of A. integerrima in the northern portion of its range, with both putative parents functioning as maternal donors, both supplying unreduced gametes, and both contributing a significant portion of their genetic diversity to the hybrids.<h4>Conclusions</h4>Our results highlight the extreme cryptic genetic diversity and systematic complexity that can underlie a single polyploid taxon.}, Doi = {10.3732/ajb.1200199}, Key = {fds230106} } @article{fds230108, Author = {Johnson, AK and Rothfels, CJ and Windham, MD and Pryer, KM}, Title = {Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae).}, Journal = {American journal of botany}, Volume = {99}, Number = {6}, Pages = {1118-1124}, Year = {2012}, Month = {June}, url = {http://www.ncbi.nlm.nih.gov/pubmed/22542903}, Abstract = {<h4>Premise of the study</h4>Not all ferns grow in moist, shaded habitats; some lineages thrive in exposed, seasonally dry environments. Notholaenids are a clade of xeric-adapted ferns commonly characterized by the presence of a waxy exudate, called farina, on the undersides of their leaves. Although some other lineages of cheilanthoid ferns also have farinose sporophytes, previous studies suggested that notholaenids are unique in also producing farina on their gametophytes. For this reason, consistent farina expression across life cycle phases has been proposed as a potential synapomorphy for the genus Notholaena. Recent phylogenetic studies have shown two species with nonfarinose sporophytes to be nested within Notholaena, with a third nonfarinose species well supported as sister to all other notholaenids. This finding raises the question: are the gametophytes of these three species farinose like those of their close relatives, or are they glabrous, consistent with their sporophytes?<h4>Methods</h4>We sowed spores of a diversity of cheilanthoid ferns onto culture media to observe and document whether their gametophytes produced farina. To place these species within a phylogenetic context, we extracted genomic DNA, then amplified and sequenced three plastid loci. The aligned data were analyzed using maximum likelihood to generate a phylogenetic tree.<h4>Key results</h4>Here we show that notholaenids lacking sporophytic farina also lack farina in the gametophytic phase, and notholaenids with sporophytic farina always display gametophytic farina (with a single exception). Outgroup taxa never displayed gametophytic farina, regardless of whether they displayed farina on their sporophytes.<h4>Conclusions</h4>Notholaenids are unique among ferns in consistently expressing farina across both phases of the life cycle.}, Doi = {10.3732/ajb.1200049}, Key = {fds230108} } @article{fds230112, Author = {Rothfels, CJ and Larsson, A and Kuo, L-Y and Korall, P and Chiou, W-L and Pryer, KM}, Title = {Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns.}, Journal = {Systematic biology}, Volume = {61}, Number = {3}, Pages = {490-509}, Year = {2012}, Month = {May}, url = {http://www.ncbi.nlm.nih.gov/pubmed/22223449}, Abstract = {Backbone relationships within the large eupolypod II clade, which includes nearly a third of extant fern species, have resisted elucidation by both molecular and morphological data. Earlier studies suggest that much of the phylogenetic intractability of this group is due to three factors: (i) a long root that reduces apparent levels of support in the ingroup; (ii) long ingroup branches subtended by a series of very short backbone internodes (the "ancient rapid radiation" model); and (iii) significantly heterogeneous lineage-specific rates of substitution. To resolve the eupolypod II phylogeny, with a particular emphasis on the backbone internodes, we assembled a data set of five plastid loci (atpA, atpB, matK, rbcL, and trnG-R) from a sample of 81 accessions selected to capture the deepest divergences in the clade. We then evaluated our phylogenetic hypothesis against potential confounding factors, including those induced by rooting, ancient rapid radiation, rate heterogeneity, and the Bayesian star-tree paradox artifact. While the strong support we inferred for the backbone relationships proved robust to these potential problems, their investigation revealed unexpected model-mediated impacts of outgroup composition, divergent effects of methods for countering the star-tree paradox artifact, and gave no support to concerns about the applicability of the unrooted model to data sets with heterogeneous lineage-specific rates of substitution. This study is among few to investigate these factors with empirical data, and the first to compare the performance of the two primary methods for overcoming the Bayesian star-tree paradox artifact. Among the significant phylogenetic results is the near-complete support along the eupolypod II backbone, the demonstrated paraphyly of Woodsiaceae as currently circumscribed, and the well-supported placement of the enigmatic genera Homalosorus, Diplaziopsis, and Woodsia.}, Doi = {10.1093/sysbio/sys001}, Key = {fds230112} } @article{fds230104, Author = {Pryer, KM}, Title = {The Distinguished Legacy of DMB: Donald MacPhail Britton (1923–2012)}, Journal = {American Fern Journal}, Volume = {102}, Number = {4}, Pages = {241-251}, Publisher = {The American Fern Society}, Year = {2012}, ISSN = {0002-8444}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000318196200001&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Doi = {10.1640/0002-8444-102.4.241}, Key = {fds230104} } @article{fds230107, Author = {Rothfels, CJ and Sundue, MA and Kato, M and Larsson, A and Kuo, LY and Schuettpelz, E and Pryer, KM}, Title = {A revised classification for eupolypod II ferns (Polypodiales: Polypodiopsida)}, Journal = {Taxon}, Volume = {61}, Number = {3}, Pages = {515-533}, Year = {2012}, ISSN = {0040-0262}, Abstract = {We present a family-level classification for the eupolypod II clade of leptosporangiate ferns, one of the two major lineages within the Eupolypods, and one of the few parts of the fern tree of life where family-level relationships were not well understood at the time of publication of the 2006 fern classification by Smith & al. Comprising over 2500 species, the composition and particularly the relationships among the major clades of this group have historically been contentious and defied phylogenetic resolution until very recently. Our classification reflects the most current available data, largely derived from published molecular phylogenetic studies. In comparison with the five-family (Aspleniaceae, Blechnaceae, Onocleaceae, Thelypteridaceae, Woodsiaceae) treatment of Smith & al., we recognize 10 families within the eupolypod II clade. Of these, Aspleniaceae, Thelypteridaceae, Blechnaceae, and Onocleaceae have the same composition as treated by Smith & al. Woodsiaceae, which Smith & al. acknowledged as possibly non-monophyletic in their treatment, is circumscribed here to include only Woodsia and its segregates; the other "woodsioid" taxa are divided among Athyriaceae, Cystopteridaceae, Diplaziopsidaceae, Rhachidosoraceae, and Hemidictyaceae. We provide circumscriptions for each family, which summarize their morphological, geographical, and ecological characters, as well as a dichotomous key to the eupolypod II families. Three of these families- Diplaziopsidaceae, Hemidictyaceae, and Rhachidosoraceae-were described in the past year based on molecular phylogenetic analyses; we provide here their first morphological treatment.}, Key = {fds230107} } @article{fds230111, Author = {Li, F and Pryer, KM and Windham, MD}, Title = {Gaga, a new fern genus segregated from Cheilanthes (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {37}, Number = {4}, Pages = {845-860}, Publisher = {American Society of Plant Taxonomists}, Year = {2012}, ISSN = {0363-6445}, url = {http://hdl.handle.net/10161/5988 Duke open access}, Abstract = {Ongoing molecular phylogenetic studies of cheilanthoid ferns confirm that the genus Cheilanthes (Pteridaceae) is polyphyletic. A monophyletic group of species within the hemionitid clade informally called the “C. marginata group” is here shown to be distinct from its closest relatives (the genus Aspidotis) and phylogenetically distant from the type species of Cheilanthes. This group is here segregated from Cheilanthes as the newly described genus, Gaga . In this study, we use molecular data from four DNA regions (plastid: matK, rbcL, trnG-R; and nuclear: gapCp) together with spore data to circumscribe the morphological and geographical boundaries of the new genus and investigate reticulate evolution within the group. Gaga is distinguished from Aspidotis by its rounded to attenuate (vs. mucronate) segment apices, minutely bullate margins of mature leaves (vs. smooth at 40 ×), and less prominently lustrous and striate adaxial blade surfaces. The new genus is distinguished from Cheilanthes s. s. by its strongly differentiated, inframarginal pseudoindusia, the production of 64 small or 32 large spores (vs. 32 small or 16 large) per sporangium, and usually glabrous leaf blades. A total of nineteen species are recognized within Gaga; seventeen new combinations are made, and two new species, Gaga germanotta and Gaga monstraparva , are described.}, Doi = {10.1600/036364412x656626}, Key = {fds230111} } @article{fds304307, Author = {Rothfels, CJ and Sundue, MA and Kuo, L and Larsson, A and Kato, M and Schuettpelz, E and Pryer, KM}, Title = {A revised family–level classification for eupolypod II ferns (Polypodiidae: Polypodiales)}, Journal = {Taxon}, Volume = {61}, Number = {3}, Pages = {515-533}, Year = {2012}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.1002/tax.613003}, Abstract = {We present a family-level classification for the eupolypod II clade of leptosporangiate ferns, one of the two major lineages within the Eupolypods, and one of the few parts of the fern tree of life where family-level relationships were not well understood at the time of publication of the 2006 fern classification by Smith & al. Comprising over 2500 species, the composition and particularly the relationships among the major clades of this group have historically been contentious and defied phylogenetic resolution until very recently. Our classification reflects the most current available data, largely derived from published molecular phylogenetic studies. In comparison with the five-family (Aspleniaceae, Blechnaceae, Onocleaceae, Thelypteridaceae, Woodsiaceae) treatment of Smith & al., we recognize 10 families within the eupolypod II clade. Of these, Aspleniaceae, Thelypteridaceae, Blechnaceae, and Onocleaceae have the same composition as treated by Smith & al. Woodsiaceae, which Smith & al. acknowledged as possibly non-monophyletic in their treatment, is circumscribed here to include only Woodsia and its segregates; the other "woodsioid" taxa are divided among Athyriaceae, Cystopteridaceae, Diplaziopsidaceae, Rhachidosoraceae, and Hemidictyaceae. We provide circumscriptions for each family, which summarize their morphological, geographical, and ecological characters, as well as a dichotomous key to the eupolypod II families. Three of these families- Diplaziopsidaceae, Hemidictyaceae, and Rhachidosoraceae-were described in the past year based on molecular phylogenetic analyses; we provide here their first morphological treatment.}, Doi = {10.1002/tax.613003}, Key = {fds304307} } @article{fds197465, Author = {Beck, J.B. and M.D. Windham and K.M. Pryer}, Title = {Do asexual lineages lead short evolutionary lives? A case-study from the fern genus Astrolepis}, Journal = {Evolution 65: 3217-3229.}, Year = {2011}, Key = {fds197465} } @article{fds230099, Author = {Wolf, PG and Der, JP and Duffy, AM and Davidson, JB and Grusz, AL and Pryer, KM}, Title = {The evolution of chloroplast genes and genomes in ferns}, Journal = {Plant molecular biology}, Volume = {76}, Number = {3-5}, Pages = {251-261}, Publisher = {Springer Netherlands}, Year = {2011}, url = {http://www.ncbi.nlm.nih.gov/pubmed/20976559}, Abstract = {Most of the publicly available data on chloroplast (plastid) genes and genomes come from seed plants, with relatively little information from their sister group, the ferns. Here we describe several broad evolutionary patterns and processes in fern plastid genomes (plastomes), and we include some new plastome sequence data. We review what we know about the evolutionary history of plastome structure across the fern phylogeny and we compare plastome organization and patterns of evolution in ferns to those in seed plants. A large clade of ferns is characterized by a plastome that has been reorganized with respect to the ancestral gene order (a similar order that is ancestral in seed plants). We review the sequence of inversions that gave rise to this organization. We also explore global nucleotide substitution patterns in ferns versus those found in seed plants across plastid genes, and we review the high levels of RNA editing observed in fern plastomes.}, Doi = {10.1007/s11103-010-9706-4}, Key = {fds230099} } @article{fds230100, Author = {Beck, JB and Windham, MD and Pryer, KM}, Title = {Do asexual polyploid lineages lead short evolutionary lives? A case study from the fern genus Astrolepis}, Journal = {Evolution: International Journal of Organic Evolution}, Volume = {65}, Number = {11}, Pages = {3217-3229}, Publisher = {Blackwell Publishing Inc Malden, USA}, Year = {2011}, url = {http://www.ncbi.nlm.nih.gov/pubmed/22023587}, Abstract = {A life-history transition to asexuality is typically viewed as leading to a heightened extinction risk, and a number of studies have evaluated this claim by examining the relative ages of asexual versus closely related sexual lineages. Surprisingly, a rigorous assessment of the age of an asexual plant lineage has never been published, although asexuality is extraordinarily common among plants. Here, we estimate the ages of sexual diploids and asexual polyploids in the fern genus Astrolepis using a well-supported plastid phylogeny and a relaxed-clock dating approach. The 50 asexual polyploid samples we included were conservatively estimated to comprise 19 distinct lineages, including a variety of auto- and allopolyploid genomic combinations. All were either the same age or younger than the crown group comprising their maternal sexual-diploid parents based simply on their phylogenetic position. Node ages estimated with the relaxed-clock approach indicated that the average maximum age of asexual lineages was 0.4 My, and individual lineages were on average 7 to 47 times younger than the crown- and total-ages of their sexual parents. Although the confounding association between asexuality and polyploidy precludes definite conclusions regarding the effect of asexuality, our results suggest that asexuality limits evolutionary potential in Astrolepis.}, Doi = {10.1111/j.1558-5646.2011.01362.x}, Key = {fds230100} } @article{fds230101, Author = {Sigel, EM and Windham, MD and Huiet, L and Yatskievych, G and Pryer, KM}, Title = {Species relationships and farina evolution in the cheilanthoid fern genus Argyrochosma (Pteridaceae)}, Journal = {Systematic Botany}, Volume = {36}, Number = {3}, Pages = {554-564}, Publisher = {American Society of Plant Taxonomists}, Year = {2011}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364411x583547}, Abstract = {Convergent evolution driven by adaptation to arid habitats has made it difficult to identify monophyletic taxa in the cheilanthoid ferns. Dependence on distinctive, but potentially homoplastic characters, to define major clades has resulted in a taxonomic conundrum: all of the largest cheilanthoid genera have been shown to be polyphyletic. Here we reconstruct the first comprehensive phylogeny of the strictly New World cheilanthoid genus Argyrochosma. We use our reconstruction to examine the evolution of farina (powdery leaf deposits), which has played a prominent role in the circumscription of cheilanthoid genera. Our data indicate that Argyrochosma comprises two major monophyletic groups: one exclusively non-farinose and the other primarily farinose. Within the latter group, there has been at least one evolutionary reversal (loss) of farina and the development of major chemical variants that characterize specific clades. Our phylogenetic hypothesis, in combination with spore data and chromosome counts, also provides a critical context for addressing the prevalence of polyploidy and apomixis within the genus. Evidence from these datasets provides testable hypotheses regarding reticulate evolution and suggests the presence of several previously undetected taxa of Argyrochosma.}, Doi = {10.1600/036364411x583547}, Key = {fds230101} } @article{fds230113, Author = {Li, F and Kuo, L and Rothfels, CJ and Ebihara, A and Chiou, W and Windham, MD and Pryer, KM}, Title = {rbcL and matK earn two thumbs up as the core DNA barcode for ferns}, Journal = {PLoS One}, Volume = {6}, Number = {10}, Pages = {e26597}, Publisher = {Public Library of Science}, Year = {2011}, url = {http://www.ncbi.nlm.nih.gov/pubmed/22028918}, Abstract = {<h4>Background</h4>DNA barcoding will revolutionize our understanding of fern ecology, most especially because the accurate identification of the independent but cryptic gametophyte phase of the fern's life history--an endeavor previously impossible--will finally be feasible. In this study, we assess the discriminatory power of the core plant DNA barcode (rbcL and matK), as well as alternatively proposed fern barcodes (trnH-psbA and trnL-F), across all major fern lineages. We also present plastid barcode data for two genera in the hyperdiverse polypod clade--Deparia (Woodsiaceae) and the Cheilanthes marginata group (currently being segregated as a new genus of Pteridaceae)--to further evaluate the resolving power of these loci.<h4>Principal findings</h4>Our results clearly demonstrate the value of matK data, previously unavailable in ferns because of difficulties in amplification due to a major rearrangement of the plastid genome. With its high sequence variation, matK complements rbcL to provide a two-locus barcode with strong resolving power. With sequence variation comparable to matK, trnL-F appears to be a suitable alternative barcode region in ferns, and perhaps should be added to the core barcode region if universal primer development for matK fails. In contrast, trnH-psbA shows dramatically reduced sequence variation for the majority of ferns. This is likely due to the translocation of this segment of the plastid genome into the inverted repeat regions, which are known to have a highly constrained substitution rate.<h4>Conclusions</h4>Our study provides the first endorsement of the two-locus barcode (rbcL+matK) in ferns, and favors trnL-F over trnH-psbA as a potential back-up locus. Future work should focus on gathering more fern matK sequence data to facilitate universal primer development.}, Doi = {10.1371/journal.pone.0026597}, Key = {fds230113} } @article{fds230114, Author = {Pryer, KM and Schuettpelz, E and Huiet, L and Grusz, AL and Rothfels, CJ and Avent, T and Schwartz, D and Windham, MD}, Title = {DNA barcoding exposes a case of mistaken identity in the fern horticultural trade.}, Journal = {Molecular ecology resources}, Volume = {10}, Number = {6}, Pages = {979-985}, Year = {2010}, Month = {November}, url = {http://www.ncbi.nlm.nih.gov/pubmed/21565107}, Abstract = {Using cheilanthoid ferns, we provide an example of how DNA barcoding approaches can be useful to the horticultural community for keeping plants in the trade accurately identified. We use plastid rbcL, atpA, and trnG-R sequence data to demonstrate that a fern marketed as Cheilanthes wrightii (endemic to the southwestern USA and northern Mexico) in the horticultural trade is, in fact, Cheilanthes distans (endemic to Australia and adjacent islands). Public and private (accessible with permission) databases contain a wealth of DNA sequence data that are linked to vouchered plant material. These data have uses beyond those for which they were originally generated, and they provide an important resource for fostering collaborations between the academic and horticultural communities. We strongly advocate the barcoding approach as a valuable new technology available to the horticulture industry to help correct plant identification errors in the international trade.}, Doi = {10.1111/j.1755-0998.2010.02858.x}, Key = {fds230114} } @article{fds230115, Author = {Korall, P and Schuettpelz, E and Pryer, KM}, Title = {Abrupt deceleration of molecular evolution linked to the origin of arborescence in ferns.}, Journal = {Evolution; international journal of organic evolution}, Volume = {64}, Number = {9}, Pages = {2786-2792}, Year = {2010}, Month = {September}, url = {http://www.ncbi.nlm.nih.gov/pubmed/20394660}, Abstract = {Molecular rate heterogeneity, whereby rates of molecular evolution vary among groups of organisms, is a well-documented phenomenon. Nonetheless, its causes are poorly understood. For animals, generation time is frequently cited because longer-lived species tend to have slower rates of molecular evolution than their shorter-lived counterparts. Although a similar pattern has been uncovered in flowering plants, using proxies such as growth form, the underlying process has remained elusive. Here, we find a deceleration of molecular evolutionary rate to be coupled with the origin of arborescence in ferns. Phylogenetic branch lengths within the “tree fern” clade are considerably shorter than those of closely related lineages, and our analyses demonstrate that this is due to a significant difference in molecular evolutionary rate. Reconstructions reveal that an abrupt rate deceleration coincided with the evolution of the long-lived tree-like habit at the base of the tree fern clade. This suggests that a generation time effect may well be ubiquitous across the green tree of life, and that the search for a responsible mechanism must focus on characteristics shared by all vascular plants. Discriminating among the possibilities will require contributions from various biological disciplines,but will be necessary for a full appreciation of molecular evolution.}, Doi = {10.1111/j.1558-5646.2010.01000.x}, Key = {fds230115} } @article{fds230116, Author = {Beck, JB and Windham, MD and Yatskievych, G and Pryer, KM}, Title = {A Diploids-First Approach to Species Delimitation and Interpreting Polyploid Evolution in the Fern Genus Astrolepis (Pteridaceae)}, Journal = {Systematic botany}, Volume = {35}, Number = {2}, Pages = {223-234}, Publisher = {American Society of Plant Taxonomists}, Year = {2010}, Month = {April}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364410791638388}, Abstract = {Polyploidy presents a challenge to those wishing to delimit the species within a group and reconstruct the phylogenetic relationships among these taxa. A clear understanding of the tree-like relationships among the diploid species can provide a framework upon which to reconstruct the reticulate events that gave rise to the polyploid lineages. In this study we apply this ““diploids-first”” strategy to the fern genus Astrolepis (Pteridaceae). Diploids are identified using the number of spores per sporangium and spore size. Analyses of plastid and low-copy nuclear sequence data provide well-supported estimates of phylogenetic relationships, including strong evidence for two morphologically distinctive diploid lineages not recognized in recent treatments. One of these corresponds to the type of Notholaena deltoidea, a species that has not been recognized in any modern treatment of Astrolepis. This species is resurrected here as the new combination Astrolepis deltoidea . The second novel lineage is that of a diploid initially hypothesized to exist by molecular and morphological characteristics of several established Astrolepis allopolyploids. This previously missing diploid species is described here as Astrolepis obscura.}, Doi = {10.1600/036364410791638388}, Key = {fds230116} } @article{fds353353, Author = {Larsson, A and Rothfels, CJ and Kuo, LY and Shuettpelz, E and Pryer, K and Korall, P}, Title = {Var hör hällebräken, ekbräken och svartbräknar hemma?-Äntligen en fylogeni över Eupolypods II}, Journal = {Systematikdagarna, Göteborg, 22-23 november, 2010}, Year = {2010}, Key = {fds353353} } @article{fds230074, Author = {Windham, MD and Huiet, L and Schuettpelz, E and Grusz, AL and Rothfels, C and Beck, J and Yatskievych, G and Pryer, KM}, Title = {Using Plastid and Nuclear DNA Sequences to Redraw Generic Boundaries and Demystif Species Complexes in Cheilanthoid Ferns.}, Journal = {AMERICAN FERN JOURNAL}, Volume = {99}, Number = {2}, Pages = {128-132}, Publisher = {AMER FERN SOC INC}, Year = {2009}, Month = {April}, ISSN = {0002-8444}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000269677200010&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Key = {fds230074} } @article{fds230095, Author = {Pryer, KM and Hearn, DJ}, Title = {Evolution of leaf form in marsileaceous ferns: evidence for heterochrony}, Journal = {Evolution: International Journal of Organic Evolution}, Volume = {63}, Number = {2}, Pages = {498-513}, Publisher = {Blackwell Publishing Inc Malden, USA}, Year = {2009}, url = {http://www.ncbi.nlm.nih.gov/pubmed/19154361}, Abstract = {Using an explicit phylogenetic framework, ontogenetic patterns of leaf form are compared among the three genera of marsileaceous ferns (Marsilea, Regnellidium, and Pilularia) with the outgroup Asplenium to address the hypothesis that heterochrony played a role in their evolution. We performed a Fourier analysis on a developmental sequence of leaves from individuals of these genera. Principal components analysis of the harmonic coefficients was used to characterize the ontogenetic trajectories of leaf form in a smaller dimensional space. Results of this study suggest that the "evolutionary juvenilization" observed in these leaf sequences is best described using a mixed model of heterochrony (accelerated growth rate and early termination at a simplified leaf form). The later stages of the ancestral, more complex, ontogenetic pattern were lost in Marsileaceae, giving rise to the simplified adult leaves of Marsilea, Regnellidium, and Pilularia. Life-history traits such as ephemeral and uncertain habitats, high reproductive rates, and accelerated maturation, which are typical for marsileaceous ferns, suggest that they may be "r strategists." The evidence for heterochrony presented here illustrates that it has resulted in profound ecological and morphological consequences for the entire life history of Marsileaceae.}, Doi = {10.1111/j.1558-5646.2008.00562.x}, Key = {fds230095} } @article{fds230096, Author = {Schneider, H and Smith, AR and Pryer, KM}, Title = {Is morphology really at odds with molecules in estimating fern phylogeny?}, Journal = {Systematic Botany}, Volume = {34}, Number = {3}, Pages = {455-475}, Publisher = {American Society of Plant Taxonomists}, Year = {2009}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364409789271209}, Abstract = {Using a morphological dataset of 136 vegetative and reproductive characters, we infer the tracheophyte phylogeny with an emphasis on early divergences of ferns (monilophytes). The dataset comprises morphological, anatomical, biochemical, and some DNA structural characters for a taxon sample of 35 species, including representatives of all major lineages of vascular plants, especially ferns. Phylogenetic relationships among vascular plants are reconstructed using maximum parsimony and Bayesian inference. Both approaches yield similar relationships and provide evidence for three major lineages of extant vascular plants: lycophytes, ferns, and seed plants. Lycophytes are sister to the euphyllophyte clade, which comprises the fern and seed plant lineages. The fern lineage consists of five clades: horsetails, whisk ferns, ophioglossoids, marattioids, and leptosporangiate ferns. This lineage is supported by characters of the spore wall and has a parsimony bootstrap value of 76%, although the Bayesian posterior probability is only 0.53. Each of the five fern clades is well supported, but the relationships among them lack statistical support. Our independent phylogenetic analyses of morphological evidence recover the same deep phylogenetic relationships among tracheophytes as found in previous studies utilizing DNA sequence data, but differ in some ways within seed plants and within ferns. We discuss the extensive independent evolution of the five extant fern clades and the evidence for the placement of whisk ferns and horsetails in our morphological analyses.}, Doi = {10.1600/036364409789271209}, Key = {fds230096} } @article{fds230097, Author = {Grusz, AL and Windham, MD and Pryer, KM}, Title = {Deciphering the origins of apomictic polyploids in the Cheilanthes yavapensis complex (Pteridaceae)}, Journal = {American Journal of Botany}, Volume = {96}, Number = {9}, Pages = {1636-1645}, Publisher = {Botanical Society of America}, Year = {2009}, ISSN = {0002-9122}, url = {http://www.ncbi.nlm.nih.gov/pubmed/21622350}, Abstract = {Deciphering species relationships and hybrid origins in polyploid agamic species complexes is notoriously difficult. In this study of cheilanthoid ferns, we demonstrate increased resolving power for clarifying the origins of polyploid lineages by integrating evidence from a diverse selection of biosystematic methods. The prevalence of polyploidy, hybridization, and apomixis in ferns suggests that these processes play a significant role in their evolution and diversification. Using a combination of systematic approaches, we investigated the origins of apomictic polyploids belonging to the Cheilanthes yavapensis complex. Spore studies allowed us to assess ploidy levels; plastid and nuclear DNA sequencing revealed evolutionary relationships and confirmed the putative progenitors (both maternal and paternal) of taxa of hybrid origin; enzyme electrophoretic evidence provided information on genome dosage in allopolyploids. We find here that the widespread apomictic triploid, Cheilanthes lindheimeri, is an autopolyploid derived from a rare, previously undetected sexual diploid. The apomictic triploid Cheilanthes wootonii is shown to be an interspecific hybrid between C. fendleri and C. lindheimeri, whereas the apomictic tetraploid C. yavapensis is comprised of two cryptic and geographically distinct lineages. We show that earlier morphology-based hypotheses of species relationships, while not altogether incorrect, only partially explain the complicated evolutionary history of these ferns.}, Doi = {10.3732/ajb.0900019}, Key = {fds230097} } @article{fds230098, Author = {Schuettpelz, E and Pryer, KM}, Title = {Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy}, Journal = {Proceedings of the National Academy of Sciences}, Volume = {106}, Number = {27}, Pages = {11200-11205}, Publisher = {National Academy of Sciences}, Year = {2009}, url = {http://www.ncbi.nlm.nih.gov/pubmed/19567832}, Abstract = {In today's angiosperm-dominated terrestrial ecosystems, leptosporangiate ferns are truly exceptional--accounting for 80% of the approximately 11,000 nonflowering vascular plant species. Recent studies have shown that this remarkable diversity is mostly the result of a major leptosporangiate radiation beginning in the Cretaceous, following the rise of angiosperms. This pattern is suggestive of an ecological opportunistic response, with the proliferation of flowering plants across the landscape resulting in the formation of many new niches--both on forest floors and within forest canopies--into which leptosporangiate ferns could diversify. At present, one-third of leptosporangiate species grow as epiphytes in the canopies of angiosperm-dominated tropical rain forests. However, we know too little about the evolutionary history of epiphytic ferns to assess whether or not their diversification was in fact linked to the establishment of these forests, as would be predicted by the ecological opportunistic response hypothesis. Here we provide new insight into leptosporangiate diversification and the evolution of epiphytism by integrating a 400-taxon molecular dataset with an expanded set of fossil age constraints. We find evidence for a burst of fern diversification in the Cenozoic, apparently driven by the evolution of epiphytism. Whether this explosive radiation was triggered simply by the establishment of modern angiosperm-dominated tropical rain forest canopies, or spurred on by some other large-scale extrinsic factor (e.g., climate change) remains to be determined. In either case, it is clear that in both the Cretaceous and Cenozoic, leptosporangiate ferns were adept at exploiting newly created niches in angiosperm-dominated ecosystems.}, Doi = {10.1073/pnas.0811136106}, Key = {fds230098} } @article{fds230088, Author = {Christenhusz, MJM and Tuomisto, H and Metzgar, JS and Pryer, KM}, Title = {Evolutionary relationships within the Neotropical, eusporangiate fern genus Danaea (Marattiaceae)}, Journal = {Molecular Phylogenetics and Evolution}, Volume = {46}, Number = {1}, Pages = {34-48}, Publisher = {Academic Press}, Year = {2008}, ISSN = {1055-7903}, url = {http://www.ncbi.nlm.nih.gov/pubmed/18042403}, Abstract = {Genera within the eusporangiate fern family Marattiaceae have long been neglected in taxonomic and systematic studies. Here we present the first phylogenetic hypothesis of relationships within the exclusively Neotropical genus Danaea based on a sampling of 60 specimens representing 31 species from various Neotropical sites. We used DNA sequence data from three plastid regions (atpB, rbcL, and trnL-F), morphological characters from both herbarium specimens and live plants observed in the field, and geographical and ecological information to examine evolutionary patterns. Eleven representatives of five other marattioid genera (Angiopteris, Archangiopteris, Christensenia, Macroglossum, and Marattia) were used to root the topology. We identified three well-supported clades within Danaea that are consistent with morphological characters: the "leprieurii" clade (containing species traditionally associated with the name D. elliptica), the "nodosa" clade (containing all species traditionally associated with the name D. nodosa), and the "alata" clade (containing all other species). All three clades are geographically and ecologically widely distributed, but subclades within them show various distribution patterns. Our phylogenetic hypothesis provides a robust framework within which broad questions related to the morphology, taxonomy, biogeography, evolution, and ecology of these ferns can be addressed.}, Doi = {10.1016/j.ympev.2007.09.015}, Key = {fds230088} } @article{fds230090, Author = {Nagalingum, NS and Nowak, MD and Pryer, KM}, Title = {Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia}, Journal = {Botanical Journal of the Linnean Society}, Volume = {157}, Number = {4}, Pages = {673-685}, Publisher = {Oxford University Press}, Year = {2008}, ISSN = {0024-4074}, url = {http://dx.doi.org/10.1111/j.1095-8339.2008.00806.x}, Abstract = {Heterosporous ferns (Salviniales) are a group of approximately 70 species that produce two types of spores (megaspores and microspores). Earlier broad-scale phylogenetic studies on the order typically focused on one or, at most, two species per genus. In contrast, our study samples numerous species for each genus, wherever possible, accounting for almost half of the species diversity of the order. Our analyses resolve Marsileaceae, Salviniaceae and all of the component genera as monophyletic. Salviniaceae incorporate Salvinia and Azolla; in Marsileaceae, Marsilea is sister to the clade of Regnellidium and Pilularia- this latter clade is consistently resolved, but not always strongly supported. Our individual species-level investigations for Pilularia and Salvinia, together with previously published studies on Marsilea and Azolla (Regnellidium is monotypic), provide phylogenies within all genera of heterosporous ferns. The Pilularia phylogeny reveals two groups: Group I includes the European taxa P. globulifera and P. minuta; Group II consists of P. americana, P. novae-hollandiae and P. novae-zelandiae from North America, Australia and New Zealand, respectively, and are morphologically difficult to distinguish. Based on their identical molecular sequences and morphology, we regard P. novae-hollandiae and P. novae-zelandiae to be conspecific; the name P. novae-hollandiae has nomenclatural priority. The status of P. americana requires further investigation as it consists of two geographically and genetically distinct North American groups and also shows a high degree of sequence similarity to P. novae-hollandiae. Salvinia also comprises biogeographically distinct units - a Eurasian group (S. natans and S. cucullata) and an American clade that includes the noxious weed S. molesta, as well as S. oblongifolia and S. minima.}, Doi = {10.1111/j.1095-8339.2008.00806.x}, Key = {fds230090} } @article{fds230091, Author = {Metzgar, JS and Skog, JE and Zimmer, EA and Pryer, KM}, Title = {The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci}, Journal = {Systematic Botany}, Volume = {33}, Number = {1}, Pages = {31-36}, Publisher = {American Society of Plant Taxonomists}, Year = {2008}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364408783887528}, Abstract = {To resolve phylogenetic relationships among all genera and subgenera in Osmundaceae, we analyzed over 8,500 characters of DNA sequence data from seven plastid loci (atpA, rbcL, rbcL–accD, rbcL–atpB, rps4–trnS, trnG–trnR, and trnL–trnF). Our results confirm those from earlier anatomical and single-gene (rbcL) studies that suggested Osmunda s.l. is paraphyletic. Osmunda cinnamomea is sister to the remainder of Osmundaceae (Leptopteris, Todea, and Osmunda s.s.). We support the recognition of a monotypic fourth genus, Osmundastrum, to reflect these results. We also resolve subgeneric relationships within Osmunda s.s. and find that subg. Claytosmunda is strongly supported as sister to the rest of Osmunda. A stable, well-supported classification for extant Osmundaceae is proposed, along with a key to all genera and subgenera.}, Doi = {10.1600/036364408783887528}, Key = {fds230091} } @article{fds230092, Author = {Rothfels, CJ and Windham, MD and Grusz, AL and Gastony, GJ and Pryer, KM}, Title = {Toward a monophyletic Notholaena (Pteridaceae): Resolving patterns of evolutionary convergence in xeric‐adapted ferns}, Journal = {Taxon}, Volume = {57}, Number = {3}, Pages = {712-724}, Year = {2008}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.1002/tax.573005}, Abstract = {Cheilanthoid ferns (Pteridaceae) are a diverse and ecologically important clade, unusual among ferns for their ability to colonize and diversify within xeric habitats. These extreme habitats are thought to drive the extensive evolutionary convergence, and thus morphological homoplasy, that has long thwarted a natural classification of cheilanthoid ferns. Here we present the first multigene phylogeny to focus on taxa traditionally assigned to the large genus Notholaena. New World taxa (Notholaena sensu Tryon) are only distantly related to species occurring in the Old World (Notholaena sensu Pichi Sermolli). The circumscription of Notholaena adopted in recent American floras is shown to be paraphyletic, with species usually assigned to Cheilanthes and Cheiloplecton nested within it. The position of Cheiloplecton is particularly surprising - given its well-developed false indusium and non-farinose blade, it is morphologically anomalous within the "notholaenoids". In addition to clarifying natural relationships, the phylogenetic hypothesis presented here helps to resolve outstanding nomenclatural issues and provides a basis for examining character evolution within this diverse, desert-adapted clade.}, Doi = {10.1002/tax.573005}, Key = {fds230092} } @article{fds230093, Author = {Schuettpelz, E and Grusz, AL and Windham, MD and Pryer, KM}, Title = {The utility of nuclear gapCp in resolving polyploid fern origins}, Journal = {Systematic Botany}, Volume = {33}, Number = {4}, Pages = {621-629}, Publisher = {American Society of Plant Taxonomists}, Year = {2008}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364408786500127}, Abstract = {Although polyploidy is rampant in ferns and plays a major role in shaping their diversity, the evolutionary history of many polyploid species remains poorly understood. Nuclear DNA sequences can provide valuable information for identifying polyploid origins; however, remarkably few nuclear markers have been developed specifically for ferns, and previously published primer sets do not work well in many fern lineages. In this study, we present new primer sequences for the amplification of a portion of the nuclear gapCp gene (encoding a glyceraldehyde-3-phosphate dehydrogenase). Through a broad survey across ferns, we demonstrate that these primers are nearly universal for this clade. With a case study in cheilanthoids, we show that this rapidly evolving marker is a powerful tool for discriminating between autopolyploids and allopolyploids. Our results indicate that gapCp holds considerable potential for addressing species-level questions across the fern tree of life.}, Doi = {10.1600/036364408786500127}, Key = {fds230093} } @article{fds230094, Author = {Hennequin, S and Schuettpelz, E and Pryer, KM and Ebihara, A and Dubuisson, J}, Title = {Divergence times and the evolution of epiphytism in filmy ferns (Hymenophyllaceae) revisited}, Journal = {International Journal of Plant Sciences}, Volume = {169}, Number = {9}, Pages = {1278-1287}, Publisher = {The University of Chicago Press}, Year = {2008}, ISSN = {1058-5893}, url = {http://dx.doi.org/10.1086/591983}, Abstract = {Although the phylogeny of the filmy fern family (Hymenophyllaceae) is rapidly coming into focus, much remains to be uncovered concerning the evolutionary history of this clade. In this study, we use two data sets (108-taxon rbcL+ rps4, 204-taxon rbcL) and fossil constraints to examine the diversification of filmy ferns and the evolution of their ecology within a temporal context. Our penalized likelihood analyses (with both data sets) indicate that the initial divergences within the Hymenophyllaceae (resulting in extant lineages) and those within one of the two major clades (trichomanoids) occurred in the early to middle Mesozoic. There was a considerable delay in the crown group diversification of the other major clade (hymenophylloids), which began to diversify only in the Cretaceous. Maximum likelihood and Bayesian character state reconstructions across the broadly sampled single-gene (rbcL) phylogeny do not allow us to unequivocally infer the ancestral habit for the family or for its two major clades. However, adding a second gene (rps4) with a more restricted taxon sampling results in a hypothesis in which filmy ferns were ancestrally terrestrial, with epiphytism having evolved several times independently during the Cretaceous. © 2008 by The University of Chicago. All rights reserved.}, Doi = {10.1086/591983}, Key = {fds230094} } @article{fds353355, Author = {Schuettpelz, ERIC and Pryer, KATHLEENM}, Title = {Fern phylogeny}, Journal = {Biology and evolution of ferns and lycophytes}, Volume = {395}, Pages = {416}, Publisher = {Cambridge University Press Cambridge}, Year = {2008}, Key = {fds353355} } @article{fds230082, Author = {Nagalingum, NS and Schneider, H and Pryer, KM}, Title = {Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus Marsilea}, Journal = {Systematic Botany}, Volume = {32}, Number = {1}, Pages = {16-25}, Publisher = {American Society of Plant Taxonomists}, Year = {2007}, ISSN = {0363-6445}, url = {http://dx.doi.org/10.1600/036364407780360256}, Abstract = {Using six plastid regions, we present a phylogeny for 26 species of the heterosporous fern genus Marsilea. Two well-supported groups within Marsilea are identified. Group I includes two subgroups, and is relatively species-poor. Species assignable to this group have glabrous leaves (although land leaves may have a few hairs), sporocarps lacking both a raphe and teeth, and share a preference for submerged conditions (i.e., they are intolerant of desiccation). Group II is relatively diverse, and its members have leaves that are pubescent, sporocarps that bear a raphe and from zero to two teeth, and the plants are often emergent at the edges of lakes and ponds. Within Group II, five subgroups receive robust support: three are predominantly African, one is New World, and one Old World. Phylogenetic assessment of morphological evolution suggests that the presence of an inferior sporocarp tooth and the place of sporocarp maturation are homoplastic characters, and are therefore of unreliable taxonomic use at an infrageneric level. In contrast, the presence of a raphe and superior sporocarp tooth are reliable synapomorphies for classification within Marsilea.}, Doi = {10.1600/036364407780360256}, Key = {fds230082} } @article{fds230084, Author = {Haugen, P and Bhattacharya, D and Palmer, JD and Turner, S and Lewis, LA and Pryer, KM}, Title = {Cyanobacterial ribosomal RNA genes with multiple, endonuclease-encoding group I introns}, Journal = {BMC evolutionary biology}, Volume = {7}, Number = {1}, Pages = {159}, Publisher = {BioMed Central}, Year = {2007}, url = {http://www.ncbi.nlm.nih.gov/pubmed/17825109}, Abstract = {<h4>Background</h4>Group I introns are one of the four major classes of introns as defined by their distinct splicing mechanisms. Because they catalyze their own removal from precursor transcripts, group I introns are referred to as autocatalytic introns. Group I introns are common in fungal and protist nuclear ribosomal RNA genes and in organellar genomes. In contrast, they are rare in all other organisms and genomes, including bacteria.<h4>Results</h4>Here we report five group I introns, each containing a LAGLIDADG homing endonuclease gene (HEG), in large subunit (LSU) rRNA genes of cyanobacteria. Three of the introns are located in the LSU gene of Synechococcus sp. C9, and the other two are in the LSU gene of Synechococcus lividus strain C1. Phylogenetic analyses show that these introns and their HEGs are closely related to introns and HEGs located at homologous insertion sites in organellar and bacterial rDNA genes. We also present a compilation of group I introns with homing endonuclease genes in bacteria.<h4>Conclusion</h4>We have discovered multiple HEG-containing group I introns in a single bacterial gene. To our knowledge, these are the first cases of multiple group I introns in the same bacterial gene (multiple group I introns have been reported in at least one phage gene and one prophage gene). The HEGs each contain one copy of the LAGLIDADG motif and presumably function as homodimers. Phylogenetic analysis, in conjunction with their patchy taxonomic distribution, suggests that these intron-HEG elements have been transferred horizontally among organelles and bacteria. However, the mode of transfer and the nature of the biological connections among the intron-containing organisms are unknown.}, Doi = {10.1186/1471-2148-7-159}, Key = {fds230084} } @article{fds230085, Author = {Metzgar, JS and Schneider, H and Pryer, KM}, Title = {Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae)}, Journal = {International Journal of Plant Sciences}, Volume = {168}, Number = {7}, Pages = {1045-1053}, Publisher = {The University of Chicago Press}, Year = {2007}, ISSN = {1058-5893}, url = {http://dx.doi.org/10.1086/519007}, Abstract = {A phylogeny for all extant species of the heterosporous fern genus Azolla is presented here based on more than 5000 base pairs of DNA sequence data from six plastid loci (rbcL, atpB, rps4, trnL-trnF, trnG-trnR, and rps4-trnS). Our results are in agreement with other recent molecular phylogenetic hypotheses that support the monophyly of sections Azolla and Rhizosperma and the proposed relationships within section Azolla. Divergence times are estimated within Azolla using a penalized likelihood approach, integrating data from fossils and DNA sequences. Penalized likelihood analyses estimate a divergence time of 50.7 Ma (Eocene) for the split between sections Azolla and Rhizosperma, 32.5 Ma (Oligocene) for the divergence of Azolla nilotica from A. pinnata within section Rhizosperma, and 16.3 Ma (Miocene) for the divergence of the two lineages within section Azolla (the A. rubra lineage from the A. mexicana complex).}, Doi = {10.1086/519007}, Key = {fds230085} } @article{fds230086, Author = {Korall, P and Conant, DS and Metzgar, JS and Schneider, H and Pryer, KM}, Title = {A molecular phylogeny of scaly tree ferns (Cyatheaceae)}, Journal = {American Journal of Botany}, Volume = {94}, Number = {5}, Pages = {873-886}, Publisher = {Botanical Society of America}, Year = {2007}, ISSN = {0002-9122}, url = {http://www.ncbi.nlm.nih.gov/pubmed/21636456}, Abstract = {Tree ferns recently were identified as the closest sister group to the hyperdiverse clade of ferns, the polypods. Although most of the 600 species of tree ferns are arborescent, the group encompasses a wide range of morphological variability, from diminutive members to the giant scaly tree ferns, Cyatheaceae. This well-known family comprises most of the tree fern diversity (∼500 species) and is widespread in tropical, subtropical, and south temperate regions of the world. Here we investigate the phylogenetic relationships of scaly tree ferns based on DNA sequence data from five plastid regions (rbcL, rbcL-accD IGS, rbcL-atpB IGS, trnG-trnR, and trnL-trnF). A basal dichotomy resolves Sphaeropteris as sister to all other taxa and scale features support these two clades: Sphaeropteris has conform scales, whereas all other taxa have marginate scales. The marginate-scaled clade consists of a basal trichotomy, with the three groups here termed (1) Cyathea (including Cnemidaria, Hymenophyllopsis, Trichipteris), (2) Alsophila sensu stricto, and (3) Gymnosphaera (previously recognized as a section within Alsophila) + A. capensis. Scaly tree ferns display a wide range of indusial structures, and although indusium shape is homoplastic it does contain useful phylogenetic information that supports some of the larger clades recognised.}, Doi = {10.3732/ajb.94.5.873}, Key = {fds230086} } @article{fds230087, Author = {Schuettpelz, E and Schneider, H and Huiet, L and Windham, MD and Pryer, KM}, Title = {A molecular phylogeny of the fern family Pteridaceae: assessing overall relationships and the affinities of previously unsampled genera}, Journal = {Molecular phylogenetics and evolution}, Volume = {44}, Number = {3}, Pages = {1172-1185}, Publisher = {Academic Press}, Year = {2007}, ISSN = {1055-7903}, url = {http://www.ncbi.nlm.nih.gov/pubmed/17570688}, Abstract = {The monophyletic Pteridaceae accounts for roughly 10% of extant fern diversity and occupies an unusually broad range of ecological niches, including terrestrial, epiphytic, xeric-adapted rupestral, and even aquatic species. In this study, we present the results of the first broad-scale and multi-gene phylogenetic analyses of these ferns, and determine the affinities of several previously unsampled genera. Our analyses of two newly assembled data sets (including 169 newly obtained sequences) resolve five major clades within the Pteridaceae: cryptogrammoids, ceratopteridoids, pteridoids, adiantoids, and cheilanthoids. Although the composition of these clades is in general agreement with earlier phylogenetic studies, it is very much at odds with the most recent subfamilial classification. Of the previously unsampled genera, two (Neurocallis and Ochropteris) are nested within the genus Pteris; two others (Monogramma and Rheopteris) are early diverging vittarioid ferns, with Monogramma resolved as polyphyletic; the last previously unsampled genus (Adiantopsis) occupies a rather derived position among cheilanthoids. Interestingly, some clades resolved within the Pteridaceae can be characterized by their ecological preferences, suggesting that the initial diversification in this family was tied to ecological innovation and specialization. These processes may well be the basis for the diversity and success of the Pteridaceae today.}, Doi = {10.1016/j.ympev.2007.04.011}, Key = {fds230087} } @article{fds230089, Author = {Schuettpelz, E and Pryer, KM}, Title = {Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes}, Journal = {Taxon}, Volume = {56}, Number = {4}, Pages = {1037-1050}, Publisher = {WILEY}, Year = {2007}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.2307/25065903}, Abstract = {In an effort to obtain a solid and balanced approximation of global fern phylogeny to serve as a tool for addressing large-scale evolutionary questions, we assembled and analyzed the most inclusive molecular dataset for leptosporangiate ferns to date. Three plastid genes (rbcL, atpB, atpA), totaling more than 4,000 bp, were sequenced for each of 400 leptosporangiate fern species (selected using a proportional sampling approach) and five outgroups. Maximum likelihood analysis of these data yielded an especially robust phylogeny: 80% of the nodes were supported by a maximum likelihood bootstrap percentage ≥ 70. The scope of our analysis provides unprecedented insight into overall fern relationships, not only delivering additional support for the deepest leptosporangiate divergences, but also uncovering the composition of more recently emerging clades and their relationships to one another.}, Doi = {10.2307/25065903}, Key = {fds230089} } @article{fds230076, Author = {Korall, P and Conant, DS and Schneider, H and Ueda, K and Nishida, H and Pryer, KM}, Title = {On the Phylogenetic Position of Cystodium It's Not a Tree Fern–It's a Polypod!}, Journal = {American Fern Journal}, Volume = {96}, Number = {2}, Pages = {45-53}, Publisher = {The American Fern Society}, Year = {2006}, ISSN = {0002-8444}, url = {http://dx.doi.org/10.1640/0002-8444(2006)96[45:OTPPOC]2.0.CO;2}, Abstract = {The phylogenetic position of Cystodium J. Sm. is studied here for the first time using DNA sequence data. Based on a broad sampling of leptosporangiate ferns and two plastid genes (rbcL and atpB), we show that Cystodium does not belong to the tree fern family Dicksoniaceae, as previously thought. Our results strongly support including Cystodium within the large polypod clade, and suggest its close relationship to the species-poor grade taxa at the base of the polypod topology (Sphenomeris and Lonchitis, or Saccoloma in this study). Further studies, with an expanded taxon sampling within polypods, are needed to fully understand the more precise phylogenetic relationships of Cystodium.}, Doi = {10.1640/0002-8444(2006)96[45:OTPPOC]2.0.CO;2}, Key = {fds230076} } @article{fds230078, Author = {Schuettpelz, E and Pryer, KM}, Title = {Reconciling extreme branch length differences: decoupling time and rate through the evolutionary history of filmy ferns}, Journal = {Systematic Biology}, Volume = {55}, Number = {3}, Pages = {485-502}, Publisher = {Society of Systematic Zoology}, Year = {2006}, ISSN = {1063-5157}, url = {http://www.ncbi.nlm.nih.gov/pubmed/16861211}, Abstract = {The rate of molecular evolution is not constant across the Tree of Life. Characterizing rate discrepancies and evaluating the relative roles of time and rate along branches through the past are both critical to a full understanding of evolutionary history. In this study, we explore the interactions of time and rate in filmy ferns (Hymenophyllaceae), a lineage with extreme branch length differences between the two major clades. We test for the presence of significant rate discrepancies within and between these clades, and we separate time and rate across the filmy fern phylogeny to simultaneously yield an evolutionary time scale of filmy fern diversification and reconstructions of ancestral rates of molecular evolution. Our results indicate that the branch length disparity observed between the major lineages of filmy ferns is indeed due to a significant difference in molecular evolutionary rate. The estimation of divergence times reveals that the timing of crown group diversification was not concurrent for the two lineages, and the reconstruction of ancestral rates of molecular evolution points to a substantial rate deceleration in one of the clades. Further analysis suggests that this may be due to a genome-wide deceleration in the rate of nucleotide substitution.}, Doi = {10.1080/10635150600755438}, Key = {fds230078} } @article{fds230079, Author = {Korall, P and Pryer, KM and Metzgar, JS and Schneider, H and Conant, DS}, Title = {Tree ferns: monophyletic groups and their relationships as revealed by four protein-coding plastid loci}, Journal = {Molecular Phylogenetics and Evolution}, Volume = {39}, Number = {3}, Pages = {830-845}, Publisher = {Academic Press}, Year = {2006}, ISSN = {1055-7903}, url = {http://www.ncbi.nlm.nih.gov/pubmed/16481203}, Abstract = {Tree ferns are a well-established clade within leptosporangiate ferns. Most of the 600 species (in seven families and 13 genera) are arborescent, but considerable morphological variability exists, spanning the giant scaly tree ferns (Cyatheaceae), the low, erect plants (Plagiogyriaceae), and the diminutive endemics of the Guayana Highlands (Hymenophyllopsidaceae). In this study, we investigate phylogenetic relationships within tree ferns based on analyses of four protein-coding, plastid loci (atpA, atpB, rbcL, and rps4). Our results reveal four well-supported clades, with genera of Dicksoniaceae (sensu ) interspersed among them: (A) (Loxomataceae, (Culcita, Plagiogyriaceae)), (B) (Calochlaena, (Dicksonia, Lophosoriaceae)), (C) Cibotium, and (D) Cyatheaceae, with Hymenophyllopsidaceae nested within. How these four groups are related to one other, to Thyrsopteris, or to Metaxyaceae is weakly supported. Our results show that Dicksoniaceae and Cyatheaceae, as currently recognised, are not monophyletic and new circumscriptions for these families are needed.}, Doi = {10.1016/j.ympev.2006.01.001}, Key = {fds230079} } @article{fds230080, Author = {Nagalingum, NS and Schneider, H and Pryer, KM}, Title = {Comparative morphology of reproductive structures in heterosporous water ferns and a reevaluation of the sporocarp}, Journal = {International Journal of Plant Sciences}, Volume = {167}, Number = {4}, Pages = {805-815}, Publisher = {The University of Chicago Press}, Year = {2006}, ISSN = {1058-5893}, url = {http://dx.doi.org/10.1086/503848}, Abstract = {Heterosporous water ferns (Marsileaceae and Salviniaceae) are the only extant group of plants to have evolved heterospory since the Paleozoic. These ferns possess unusual reproductive structures traditionally termed “sporocarps.” Using an evolutionary framework, we critically examine the complex homology issues pertaining to these structures. Comparative morphological study reveals that all heterosporous ferns bear indusiate sori on a branched, nonlaminate structure that we refer to as the sorophore; this expanded definition highlights homology previously obscured by the use of different terms. By using a homology-based concept, we aim to discontinue the use of historically and functionally based morphological terminology. We recognize the sorophore envelope as a structure that surrounds the sorophore and sori. The sorophore envelope is present in Marsileaceae as a sclerenchymatous sporocarp wall and in Azolla as a parenchymatous layer, but it is absent in Salvinia. Both homology assessments and phylogenetic character-state reconstructions using the Cretaceous fossil Hydropteris are consistent with a single origin of the sorophore envelope in heterosporous ferns. Consequently, we restrict the term “sporocarp” to a sorophore envelope and all it contains. Traditional usage of “sporocarp” is misleading because it implies homology for nonhomologous structures, and structures historically called sporocarps in Salviniaceae are more appropriately referred to as sori.}, Doi = {10.1086/503848}, Key = {fds230080} } @article{fds230081, Author = {Smith, AR and Pryer, KM and Schuettpelz, E and Korall, P and Schneider, H and Wolf, PG}, Title = {A classification for extant ferns}, Journal = {Taxon}, Volume = {55}, Number = {3}, Pages = {705-731}, Publisher = {WILEY}, Year = {2006}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.2307/25065646}, Abstract = {We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synopsis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11 monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family, Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including common synonyms), families, orders, and taxa of higher rank are provided.}, Doi = {10.2307/25065646}, Key = {fds230081} } @article{fds230083, Author = {Schuettpelz, E and Korall, P and Pryer, KM}, Title = {Plastid atpA data provide improved support for deep relationships among ferns}, Journal = {Taxon}, Volume = {55}, Number = {4}, Pages = {897-906}, Publisher = {WILEY}, Year = {2006}, ISSN = {0040-0262}, url = {http://dx.doi.org/10.2307/25065684}, Abstract = {DNA sequence data and phylogenetic approaches have contributed greatly to our understanding of fern relationships. Nonetheless, the datasets analyzed to date have not been sufficient to definitively resolve all parts of the global fern phylogeny; additional data and more extensive sampling are necessary. Here, we explore the phylogenetic utility of the plastid atpA gene. Using newly designed primers, we obtained atpA sequences for 52 fern and 6 outgroup taxa, and then evaluated the capabilities of atpA relative to four other molecular markers, as well as the contributions of atpA in combined analyses. The five single-gene datasets differed markedly in the number of variable characters they possessed; and although the relationships resolved in analyses of these datasets were largely congruent, the robustness of the hypotheses varied considerably. The atpA dataset had more variable characters and resulted in a more robustly supported phylogeny than any of the other single gene datasets examined, suggesting that atpA will be exceptionally useful in more extensive studies of fern phylogeny and perhaps also in studies of other plant lineages. When the atpA data were analyzed in combination with the other four markers, an especially robust hypothesis of fern relationships emerged. With the addition of the atpA data, support increased substantially at several nodes; three nodes, which were not well-supported previously, received both good posterior probability and good bootstrap support in the combined 5-gene (> 6 kb) analyses.}, Doi = {10.2307/25065684}, Key = {fds230083} } @article{fds230077, Author = {Wikström, N and Pryer, KM}, Title = {Incongruence between primary sequence data and the distribution of a mitochondrial atp1 group II intron among ferns and horsetails}, Journal = {Molecular Phylogenetics and Evolution}, Volume = {36}, Number = {3}, Pages = {484-493}, Publisher = {Academic Press}, Year = {2005}, ISSN = {1055-7903}, url = {http://www.ncbi.nlm.nih.gov/pubmed/15922630}, Abstract = {Using DNA sequence data from multiple genes (often from more than one genome compartment) to reconstruct phylogenetic relationships has become routine. Augmenting this approach with genomic structural characters (e.g., intron gain and loss, changes in gene order) as these data become available from comparative studies already has provided critical insight into some long-standing questions about the evolution of land plants. Here we report on the presence of a group II intron located in the mitochondrial atp1 gene of leptosporangiate and marattioid ferns. Primary sequence data for the atp1 gene are newly reported for 27 taxa, and results are presented from maximum likelihood-based phylogenetic analyses using Bayesian inference for 34 land plants in three data sets: (1) single-gene mitochondrial atp1 (exon+intron sequences); (2) five combined genes (mitochondrial atp1 [exon only]; plastid rbcL, atpB, rps4; nuclear SSU rDNA); and (3) same five combined genes plus morphology. All our phylogenetic analyses corroborate results from previous fern studies that used plastid and nuclear sequence data: the monophyly of euphyllophytes, as well as of monilophytes; whisk ferns (Psilotidae) sister to ophioglossoid ferns (Ophioglossidae); horsetails (Equisetopsida) sister to marattioid ferns (Marattiidae), which together are sister to the monophyletic leptosporangiate ferns. In contrast to the results from the primary sequence data, the genomic structural data (atp1 intron distribution pattern) would seem to suggest that leptosporangiate and marattioid ferns are monophyletic, and together they are the sister group to horsetails--a topology that is rarely reconstructed using primary sequence data.}, Doi = {10.1016/j.ympev.2005.04.008}, Key = {fds230077} } @article{fds230118, Author = {Pryer, KM and Schuettpelz, E and Wolf, PG and Schneider, H and Smith, AR and Cranfill, R}, Title = {Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences.}, Journal = {American journal of botany}, Volume = {91}, Number = {10}, Pages = {1582-1598}, Year = {2004}, Month = {October}, ISSN = {0002-9122}, url = {http://www.ncbi.nlm.nih.gov/pubmed/21652310}, Abstract = {The phylogenetic structure of ferns (= monilophytes) is explored here, with a special focus on the early divergences among leptosporangiate lineages. Despite considerable progress in our understanding of fern relationships, a rigorous and comprehensive analysis of the early leptosporangiate divergences was lacking. Therefore, a data set was designed here to include critical taxa that were not included in earlier studies. More than 5000 bp from the plastid (rbcL, atpB, rps4) and the nuclear (18S rDNA) genomes were sequenced for 62 taxa. Phylogenetic analyses of these data (1) confirm that Osmundaceae are sister to the rest of the leptosporangiates, (2) resolve a diverse set of ferns formerly thought to be a subsequent grade as possibly monophyletic (((Dipteridaceae, Matoniaceae), Gleicheniaceae), Hymenophyllaceae), and (3) place schizaeoid ferns as sister to a large clade of "core leptosporangiates" that includes heterosporous ferns, tree ferns, and polypods. Divergence time estimates for ferns are reported from penalized likelihood analyses of our molecular data, with constraints from a reassessment of the fossil record.}, Doi = {10.3732/ajb.91.10.1582}, Key = {fds230118} } @article{fds230117, Author = {Schneider, H and Schuettpelz, E and Pryer, KM and Cranfill, R and Magallón, S and Lupia, R}, Title = {Ferns diversified in the shadow of angiosperms.}, Journal = {Nature}, Volume = {428}, Number = {6982}, Pages = {553-557}, Year = {2004}, Month = {April}, url = {http://www.ncbi.nlm.nih.gov/pubmed/15058303}, Abstract = {The rise of angiosperms during the Cretaceous period is often portrayed as coincident with a dramatic drop in the diversity and abundance of many seed-free vascular plant lineages, including ferns. This has led to the widespread belief that ferns, once a principal component of terrestrial ecosystems, succumbed to the ecological predominance of angiosperms and are mostly evolutionary holdovers from the late Palaeozoic/early Mesozoic era. The first appearance of many modern fern genera in the early Tertiary fossil record implies another evolutionary scenario; that is, that the majority of living ferns resulted from a more recent diversification. But a full understanding of trends in fern diversification and evolution using only palaeobotanical evidence is hindered by the poor taxonomic resolution of the fern fossil record in the Cretaceous. Here we report divergence time estimates for ferns and angiosperms based on molecular data, with constraints from a reassessment of the fossil record. We show that polypod ferns (> 80% of living fern species) diversified in the Cretaceous, after angiosperms, suggesting perhaps an ecological opportunistic response to the diversification of angiosperms, as angiosperms came to dominate terrestrial ecosystems.}, Doi = {10.1038/nature02361}, Key = {fds230117} } @article{fds353357, Author = {LUPIA, R and NOWAK, M and SCHNEIDER, H and NAGALINGUM, N and PRYER, K}, Title = {HETEROSPOROUS FERNS: PHYLOGENY, ORIGIN AND RADIATION}, Journal = {2004 Denver Annual Meeting}, Year = {2004}, Key = {fds353357} } @article{fds230120, Author = {Des Marais and DL and Smith, AR and Britton, DM and Pryer, KM}, Title = {Phylogenetic relationships and evolution of extant Horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F).}, Journal = {International journal of plant sciences}, Volume = {164}, Number = {5}, Pages = {737-751}, Publisher = {University of Chicago Press}, Year = {2003}, Month = {September}, url = {http://dx.doi.org/10.1086/376817}, Abstract = {Equisetum is a small and morphologically distinct genus with a rich fossil record. Two subgenera have been recognized based principally on stomatal position and stem branching: subg. Equisetum (eight species; superficial stomates; stems branched) and subg. Hippochaete (seven species; sunken stomates; stems generally unbranched). Prior attempts at understanding Equisetum systematics, phylogeny, and character evolution have been hampered by the high degree of morphological plasticity in the genus as well as by frequent hybridization among members within each subgenus. We present the first explicit phylogenetic study of Equisetum, including all 15 species and two samples of one widespread hybrid, Equisetum ×ferrissii, based on a combined analysis of two chloroplast markers, rbcL and trnL-F. Our robustly supported phylogeny identifies two monophyletic clades corresponding to the two subgenera recognized by earlier workers. The phylogenetic placement of Equisetum bogotense, however, is ambiguous. In maximum likelihood analyses, it allies with subg. Hippochaete as the most basal member, while maximum parsimony places it as sister to the rest of the genus. A consensus phylogeny from the two analyses is presented as a basal trichotomy (E. bogotense, subg. Hippochaete, subg. Equisetum), and morphological character evolution is discussed. We detected rate heterogeneity in the rbcL locus between the two subgenera that can be attributed to an increased rate of nucleotide substitution (transversions) in subg. Hippochaete. We calculated molecular-based age estimates using the penalized likelihood approach, which accounts for rate heterogeneity and does not assume a molecular clock. The Equisetum crown group appears to have diversified in the early Cenozoic, whereas the Equisetaceae total group is estimated to have a Paleozoic origin. These molecular-based age estimates are in remarkable agreement with current interpretations of the fossil record.}, Doi = {10.1086/376817}, Key = {fds230120} } @article{fds230121, Author = {Dubuisson, JY and Hennequin, S and Douzery, EJP and Cranfill, RB and Smith, AR and Pryer, KM}, Title = {rbcL phylogeny of the fern genus Trichomanes (Hymenophyllaceae), with special reference to neotropical taxa.}, Journal = {International journal of plant sciences}, Volume = {164}, Number = {5}, Pages = {753-761}, Publisher = {University of Chicago Press}, Year = {2003}, Month = {September}, url = {http://dx.doi.org/10.1086/377059}, Abstract = {In order to estimate evolutionary relationships within the filmy fern genus Trichomanes (Hymenophyllaceae), we performed a phylogenetic analysis using rbcL nucleotide data from 46 species of Trichomanes belonging to all four of C. V. Morton's subgenera: Achomanes, Didymoglossum, Pachychaetum, and Trichomanes. Outgroups included four species of Hymenophyllum in three different subgenera, plus the monotypic genus Cardiomanes, from New Zealand. We find high resolution and robust support at most nodes, regardless of the phylogenetic optimization criterion used (maximum parsimony or maximum likelihood). Two species belonging to Morton's Asiatic sections Callistopteris and Cephalomanes are in unresolved basal positions within Trichomanes s.l., suggesting that rbcL data alone are inadequate for estimating the earliest cladogenetic events. Out of the four Morton trichomanoid subgenera, only subg. Didymoglossum appears monophyletic. Other noteworthy results include the following: (1) lianescent sect. Lacostea is more closely related to sect. Davalliopsis (traditionally placed in subg. Pachychaetum) than to other members of subg. Achomanes; (2) sections Davalliopsis and Lacostea, together with species of the morphologically different subg. Achomanes, make up a strongly supported Neotropical clade; (3) all hemiepiphytes (but not true lianas) and strictly epiphytic or epipetric species (Morton's subgenera Trichomanes and Didymoglossum) group together in an ecologically definable clade that also includes the terrestrial sect. Nesopteris; and (4) sect. Lacosteopsis (sensu Morton) is polyphyletic and comprises two distantly related clades: large hemiepiphytic climbers and small strictly epiphytic/epipetric taxa. Each of these associations is somewhat unexpected but is supported by cytological, geographical, and/or ecological evidence. We conclude that many morphological characters traditionally used for delimiting groups within Trichomanes are, in part, plesiomorphic or homoplastic. Additionally, we discuss probable multiple origins of Neotropical Trichomanes.}, Doi = {10.1086/377059}, Key = {fds230121} } @article{fds353358, Author = {Pryer, KM and Schuettpelz, E and Cranfill, RC and Wolf, PG and Smith, AR and Schneider, H}, Title = {Phylogeny of Early-Diverging Leptosporangiate Ferns Based on Morphology and Multiple Genes: rbcL, atpB, rps4, and 18S}, Year = {2003}, Key = {fds353358} } @article{fds230122, Author = {Pryer, KM and Schneider, H and Zimmer, EA and Ann Banks, J}, Title = {Deciding among green plants for whole genome studies.}, Journal = {Trends in plant science}, Volume = {7}, Number = {12}, Pages = {550-554}, Year = {2002}, Month = {December}, ISSN = {1360-1385}, url = {http://www.ncbi.nlm.nih.gov/pubmed/12475497}, Abstract = {Recent comparative DNA-sequencing studies of chloroplast, mitochondrial and ribosomal genes have produced an evolutionary tree relating the diversity of green-plant lineages. By coupling this phylogenetic framework to the explosion of information on genome content, plant-genomic efforts can and should be extended beyond angiosperm crop and model systems. Including plant species representative of other crucial evolutionary nodes would produce the comparative information necessary to understand fully the organization, function and evolution of plant genomes. The simultaneous development of genomic tools for green algae, bryophytes, 'seed-free' vascular plants and gymnosperms should provide insights into the bases of the complex morphological, physiological, reproductive and biochemical innovations that have characterized the successful transition of green plants to land.}, Doi = {10.1016/s1360-1385(02)02375-0}, Key = {fds230122} } @article{fds230123, Author = {Schneider, H and Pryer, KM}, Title = {Structure and function of spores in the aquatic heterosporous fern family Marsileaceae.}, Journal = {International journal of plant sciences}, Volume = {163}, Number = {4}, Pages = {485-505}, Publisher = {University of Chicago Press}, Year = {2002}, Month = {July}, url = {http://dx.doi.org/10.1086/340736}, Abstract = {Spores of the aquatic heterosporous fern family Marsileaceae differ markedly from spores of Salviniaceae, the only other family of heterosporous ferns and sister group to Marsileaceae, and from spores of all homosporous ferns. The marsileaceous outer spore wall (perine) is modified above the aperture into a structure, the acrolamella, and the perine and acrolamella are further modified into a remarkable gelatinous layer that envelops the spore. Observations with light and scanning electron microscopy indicate that the three living marsileaceous fern genera (Marsilea, Pilularia, and Regnellidium) each have distinctive spores, particularly with regard to the perine and acrolamella. Several spore characters support a division of Marsilea into two groups. Spore character evolution is discussed in the context of developmental and possible functional aspects. The gelatinous perine layer acts as a flexible, floating organ that envelops the spores only for a short time and appears to be an adaptation of marsileaceous ferns to amphibious habitats. The gelatinous nature of the perine layer is likely the result of acidic polysaccharide components in the spore wall that have hydrogel (swelling and shrinking) properties. Megaspores floating at the water/air interface form a concave meniscus, at the center of which is the gelatinous acrolamella that encloses a “sperm lake.” This meniscus creates a vortex-like effect that serves as a trap for free-swimming sperm cells, propelling them into the sperm lake.}, Doi = {10.1086/340736}, Key = {fds230123} } @article{fds230124, Author = {Smith, AR and Tuomisto, H and Pryer, KM and Hunt, JS and Wolf, PG}, Title = {Metaxya lanosa, a second species in the genus and fern family Metaxyaceae}, Journal = {Systematic Botany}, Volume = {26}, Number = {3}, Pages = {480-486}, Year = {2001}, Month = {October}, ISSN = {0363-6445}, Abstract = {We describe and illustrate Metaxya lanosa, the second known species in the genus and the fern family Metaxyaceae (Pteridophyta). It is currently known from four different watersheds in Amazonian Peru and Venezuela. It can be distinguished readily from M. rostrata by the noticeably woolly-hairy stipes and rachises (hairs red-brown or orange-brown and easily abraded), broader, more elliptic pinnae, cartilaginous and whitish pinna margins, more distinct veins abaxially, and longer pinna stalks, especially on the distal pinnae, rbcL data from a very limited sampling are ambiguous but do not reject support for the recognition of at least two species within Metaxya.}, Key = {fds230124} } @article{fds230125, Author = {Pryer, KM and Smith, AR and Hunt, JS and Dubuisson, JY}, Title = {rbcL data reveal two monophyletic groups of filmy ferns (Filicopsida: Hymenophyllaceae).}, Journal = {American journal of botany}, Volume = {88}, Number = {6}, Pages = {1118-1130}, Year = {2001}, Month = {June}, ISSN = {0002-9122}, url = {http://www.ncbi.nlm.nih.gov/pubmed/11410477}, Abstract = {The "filmy fern" family, Hymenophyllaceae, is traditionally partitioned into two principal genera, Trichomanes s.l. (sensu lato) and Hymenophyllum s.l., based upon sorus shape characters. This basic split in the family has been widely debated this past century and hence was evaluated here by using rbcL nucleotide sequence data in a phylogenetic study of 26 filmy ferns and nine outgroup taxa. Our results confirm the monophyly of the family and provide robust support for two monophyletic groups that correspond to the two classical genera. In addition, we show that some taxa of uncertain affinity, such as the monotypic genera Cardiomanes and Serpyllopsis, and at least one species of Microtrichomanes, are convincingly included within Hymenophyllum s.l. The tubular- or conical-based sorus that typifies Trichomanes s.l. and Cardiomanes, the most basal member of Hymenophyllum s.l., is a plesiomorphic character state for the family. Tubular-based sori occurring in other members of Hymenophyllum s.l. are most likely derived independently and more than one time. While rbcL data are able to provide a well-supported phylogenetic estimate within Trichomanes s.l., they are inadequate for resolving relationships within Hymenophyllum s.l., which will require data from additional sources. This disparity in resolution reflects differential rates of evolution for rbcL within Hymenophyllaceae.}, Doi = {10.2307/2657095}, Key = {fds230125} } @article{fds230126, Author = {Pryer, KM and Schneider, H and Smith, AR and Cranfill, R and Wolf, PG and Hunt, JS and Sipes, SD}, Title = {Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants.}, Journal = {Nature}, Volume = {409}, Number = {6820}, Pages = {618-622}, Year = {2001}, Month = {February}, ISSN = {0028-0836}, url = {http://www.ncbi.nlm.nih.gov/pubmed/11214320}, Abstract = {Most of the 470-million-year history of plants on land belongs to bryophytes, pteridophytes and gymnosperms, which eventually yielded to the ecological dominance by angiosperms 90 Myr ago. Our knowledge of angiosperm phylogeny, particularly the branching order of the earliest lineages, has recently been increased by the concurrence of multigene sequence analyses. However, reconstructing relationships for all the main lineages of vascular plants that diverged since the Devonian period has remained a challenge. Here we report phylogenetic analyses of combined data--from morphology and from four genes--for 35 representatives from all the main lineages of land plants. We show that there are three monophyletic groups of extant vascular plants: (1) lycophytes, (2) seed plants and (3) a clade including equisetophytes (horsetails), psilotophytes (whisk ferns) and all eusporangiate and leptosporangiate ferns. Our maximum-likelihood analysis shows unambiguously that horsetails and ferns together are the closest relatives to seed plants. This refutes the prevailing view that horsetails and ferns are transitional evolutionary grades between bryophytes and seed plants, and has important implications for our understanding of the development and evolution of plants.}, Doi = {10.1038/35054555}, Key = {fds230126} } @article{fds353360, Author = {Pryer, KM and Schneider, H and Smith, AR and Wolf, PG}, Title = {The Closest Living Relative to Seed Plants: Insights from Four Genes and Morphology}, Volume = {87 (Suppl)}, Pages = {151}, Year = {2001}, Key = {fds353360} } @article{fds230119, Author = {Lupia, R and Schneider, H and Moeser, GM and Pryer, KM and Crane, PR}, Title = {Marsileaceae sporocarps and spores from the late cretaceous of Georgia, U.S.A.}, Journal = {International Journal of Plant Sciences}, Volume = {161}, Number = {6}, Pages = {975-988}, Publisher = {University of Chicago Press}, Year = {2000}, Month = {January}, url = {http://dx.doi.org/10.1086/317567}, Abstract = {A new species provisionally assigned to the extant genus Regnellidium Lindm. (Regnellidium upatoiensis sp. nov.) is established for isolated sporocarps assignable to the heterosporous water fern family Marsileaceae. Three sporocarps and hundreds of dispersed megaspores were recovered from unconsolidated clays and silts of the Eutaw Formation (Santonian, Late Cretaceous) along Upatoi Creek, Georgia, U.S.A. The sporocarps are ellipsoidal and flattened, contain both megasporangia and microsporangia, and possess a two-layered wall - an outer sclerenchymatous layer and an inner parenchymatous layer. In situ megaspores are spheroidal, with two distinct wall layers - an exine, differentiated into two layers, and an outer ornamented perine also differentiated into two layers. The megaspores also possess an acrolamella consisting of six (five to seven) triangular lobes that are twisted. In situ microspores are trilete and spheroidal, with a strongly rugulate perine, and show modification of the perine over the laesura to form an acrolamella. Comparison of the fossil sporocarps with those of four extant species of Marsileaceae reveal marked similarity with Regnellidium diphyllum Lindm., particularly in megaspore and microspore morphology. If found dispersed, the in situ megaspores would be assigned to Molaspora lobata (Dijkstra) Hall and the microspores to Crybelosporites Dettmann based on their size, shape, and ornamentation. Regnellidium upatoiensis sp. nov. extends the stratigraphic range of the genus back to the Santonian, nearly contemporaneous with the first evidence of Marsilea, and implies that the diversification of the Marsileaceae into its extant lineages occurred in the mid-Cretaceous.}, Doi = {10.1086/317567}, Key = {fds230119} } @article{fds353361, Author = {Schneider, H and Pryer, KM and Smith, AR and Wolf, PG and Cranfill, RC}, Title = {Phylogeny of vascular plants and the evolution of their body plans}, Year = {2000}, Key = {fds353361} } @article{fds230133, Author = {Pryer, KM}, Title = {Phylogeny of Marsileaceous Ferns and Relationships of the Fossil Hydropteris pinnata Reconsidered.}, Journal = {International journal of plant sciences}, Volume = {160}, Number = {5}, Pages = {931-954}, Year = {1999}, Month = {September}, ISSN = {1058-5893}, url = {http://www.ncbi.nlm.nih.gov/pubmed/10506474}, Abstract = {Recent phylogenetic studies have provided compelling evidence that confirms the once disputed hypothesis of monophyly for heterosporous leptosporangiate ferns (Marsileaceae and Salviniaceae). Hypotheses for relationships among the three genera of Marsileaceae (Marsilea, Regnellidium, and Pilularia), however, have continued to be in conflict. The phylogeny of Marsileaceae is investigated here using information from morphology and rbcL sequence data. In addition, relationships among all heterosporous ferns, including the whole-plant fossil Hydropteris pinnata are reconsidered. Data sets of 71 morphological and 1239 rbcL characters for 23 leptosporangiate ferns, including eight heterosporous ingroup taxa and 15 homosporous outgroup taxa, were subjected to maximum parsimony analysis. Morphological analyses were carried out both with and without the fossil Hydropteris, and it was excluded from all analyses with rbcL data. An annotated list of the 71 morphological characters is provided in the appendix. For comparative purposes, the Rothwell and Stockey (1994) data set was also reanalyzed here. The best estimate of phylogenetic relationships for Marsileaceae in all analyses is that Pilularia and Regnellidium are sister taxa and Marsilea is sister to that clade. Morphological synapomorphies for various nodes are discussed. Analyses that included Hydropteris resulted in two most-parsimonious trees that differ only in the placement of the fossil. One topology is identical to the relationship found by Rothwell and Stockey (1994), placing the fossil sister to the Azolla plus Salvinia clade. The alternative topology places Hydropteris as the most basal member of the heterosporous fern clade. Equivocal interpretations for character evolution in heterosporous ferns are discussed in the context of these two most-parsimonious trees. Because of the observed degree of character ambiguity, the phylogenetic placement of Hydropteris is best viewed as unresolved, and recognition of the suborder Hydropteridineae, as circumscribed by Rothwell and Stockey (1994), is regarded as premature. The two competing hypotheses of relationships for heterosporous ferns are also compared with the known temporal distribution of relevant taxa. Stratigraphic fit of the phylogenetic estimates is measured by using the Stratigraphic Consistency Index and by comparison with minimum divergence times.}, Doi = {10.1086/314177}, Key = {fds230133} } @article{fds230140, Author = {Turner, S and Pryer, KM and Miao, VP and Palmer, JD}, Title = {Investigating deep phylogenetic relationships among cyanobacteria and plastids by small subunit rRNA sequence analysis.}, Journal = {The Journal of eukaryotic microbiology}, Volume = {46}, Number = {4}, Pages = {327-338}, Publisher = {Lawrence, Kan.: Society of Protozoologists, c1993-}, Year = {1999}, Month = {July}, ISSN = {1066-5234}, url = {http://www.ncbi.nlm.nih.gov/pubmed/10461381}, Abstract = {Small subunit rRNA sequence data were generated for 27 strains of cyanobacteria and incorporated into a phylogenetic analysis of 1,377 aligned sequence positions from a diverse sampling of 53 cyanobacteria and 10 photosynthetic plastids. Tree inference was carried out using a maximum likelihood method with correction for site-to-site variation in evolutionary rate. Confidence in the inferred phylogenetic relationships was determined by construction of a majority-rule consensus tree based on alternative topologies not considered to be statistically significantly different from the optimal tree. The results are in agreement with earlier studies in the assignment of individual taxa to specific sequence groups. Several relationships not previously noted among sequence groups are indicated, whereas other relationships previously supported are contradicted. All plastids cluster as a strongly supported monophyletic group arising near the root of the cyanobacterial line of descent.}, Doi = {10.1111/j.1550-7408.1999.tb04612.x}, Key = {fds230140} } @article{fds230129, Author = {Wolf, PG and Sipes, SD and White, MR and Martines, ML and Pryer, KM and Smith, AR and Ueda, K}, Title = {Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: Evidence from rbcL nucleotide sequences}, Journal = {Plant Systematics and Evolution}, Volume = {219}, Number = {3-4}, Pages = {263-270}, Publisher = {Springer Nature}, Year = {1999}, Month = {January}, url = {http://dx.doi.org/10.1007/BF00985583}, Abstract = {Nucleotide sequences from rbcL were used to infer relationships of Lophosoriaceae and Hymenophyllopsidaceae. The phylogenetic positions of these two monotypic fern families have been debated, and neither group had been included in recent molecular systematic studies of ferns. Maximum parsimony analysis of our data supported a sister relationship between Lophosoria and Dicksonia, and also between Hymenophyllopsis and Cyathea. Thus, both newly-examined families appear to be part of a previously characterized and well-supported clade of tree ferns. The inferred relationships of Lophosoria are consistent with most (but not all) recent treatments. However, Hymenophyllopsis includes only small delicate plants superficially similar to filmy ferns (Hymenophyllaceae), very different from the large arborescent taxa. Nevertheless, some synapomorphic characteristics are shared with the tree fern clade. Further studies on gametophytes of Hymenophyllopsis are needed to test these hypotheses of relationship.}, Doi = {10.1007/BF00985583}, Key = {fds230129} } @article{fds16716, Author = {Pryer, K.M. and S.J. Hackett}, Title = {Field’s Museum’s Pritzker Laboratory of Molecular Systematics and Evolution}, Year = {1999}, url = {http://www.fmnh.org/research_collections/pritzker_lab/pritzker/index.html}, Key = {fds16716} } @article{fds16717, Author = {Pryer, K.M.}, Title = {Phylogeny, character evolution, and diversification of extant ferns}, Year = {1999}, url = {http://www.fmnh.org/research_collections/botany/botany_sites/ferns/index.html}, Key = {fds16717} } @article{fds353363, Author = {Pryer, KM and Schneider, H and Hunt, JS and Wolf, PG and Smith, AR}, Title = {Basal Tracheophytes and the Phylogeny of" Pteridophyte" Lineages}, Year = {1999}, Key = {fds353363} } @article{fds353365, Author = {Hunt, JS and Pryer, KM and Vaghani, A and Smith, AR and Wolf, PG}, Title = {Fern DNA Database: Using Filemaker Pro to Coordinate DNA-Availability, DNA-Sequence Data, Vaoucher and Source Information for Large-Scale and Collaborative Phylogeny Studies}, Year = {1999}, Key = {fds353365} } @article{fds353366, Author = {Pryer, KM and Smith, AR}, Title = {Leptosporangiate ferns}, Year = {1998}, url = {http://phylogeny.arizona.edu/tree/eukaryotes/green_plants/}, Abstract = {The Tree of Life: A distributed Internet project containing information about phylogeny and biodiversity.}, Key = {fds353366} } @article{fds16719, Author = {Pryer, K.M. and A.R. Smith}, Title = {Filicopsida}, Journal = {Maddison, D.R. and W. P. Maddison}, Year = {1997}, url = {http://phylogeny.arizona.edu/tree/eukaryotes/green_plants/}, Abstract = {The Tree of Life: A distributed Internet project containing information about phylogeny and biodiversity.}, Key = {fds16719} } @article{fds353367, Author = {Pryer, KM}, Title = {Phylogenetic relationships of extant ferns, with a special emphasis on the Marsileaceae.}, Year = {1997}, Key = {fds353367} } @article{fds230127, Author = {Pryer, KM and Smith, AR and Skog, JE}, Title = {Phylogenetic Relationships of Extant Ferns Based on Evidence from Morphology and rbcL Sequences}, Journal = {American Fern Journal}, Volume = {85}, Number = {4}, Pages = {205-205}, Publisher = {JSTOR}, Year = {1995}, Month = {October}, ISSN = {0002-8444}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1995TY07900007&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Doi = {10.2307/1547810}, Key = {fds230127} } @article{fds230128, Author = {Hasebe, M and Wolf, PG and Pryer, KM and Ueda, K and Ito, M and Sano, R and Gastony, GJ and Yokoyama, J and Manhart, JR and Murakami, N and Crane, EH and Haufler, CH and Hauk, WD}, Title = {Fern Phylogeny Based on rbcL Nucleotide Sequences}, Journal = {American Fern Journal}, Volume = {85}, Number = {4}, Pages = {134-134}, Publisher = {JSTOR}, Year = {1995}, Month = {October}, ISSN = {0002-8444}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1995TY07900004&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Doi = {10.2307/1547807}, Key = {fds230128} } @article{fds230130, Author = {Pryer, KM and Haufler, CH}, Title = {Isozymic and chromosomal evidence for the allotetraploid origin of Gymnocarpium dryopteris (Dryopteridaceae)}, Journal = {Systematic botany}, Volume = {18}, Number = {1}, Pages = {150-172}, Publisher = {JSTOR}, Year = {1993}, Month = {January}, ISSN = {0363-6445}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1993KH85000013&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Abstract = {A critical examination of isozyme, chromosomal, and morphological characters in subspecies formerly included in Gymnocarpium dryopteris demonstrated that three sexual taxa can be distinguished. We recognize these taxa as distinct species: the widespread, fertile allotetraploid G. dryopteris, with one genome derived from the western diploid G. disjunctum and the other from G. appalachianum sp. nov., a previously undetected eastern North American diploid, which is described and illustrated here and for which we report a chromosome number of 2n = 80. Population comparisons of allele frequencies between C. disjunctum and G. appalachianum yielded an average Nei's genetic identity value (I) of 0.274. A wide-ranging assemblage of putatively triploid plants with both sterile, malformed spores and large, round spores capable of germination is believed to represent the backcrosses G. disjunctum X dryopteris and G. appalachianum X dryopteris. The name G. X brittonianum comb. nov. is applied here to G. disjunctum X dryopteris. A key to fertile species, species descriptions and illustrations, distribution maps, and habitat notes are included.}, Doi = {10.2307/2419795}, Key = {fds230130} } @article{fds16676, Author = {Pryer, K.M.}, Title = {Gymnocarpium}, Series = {Volume 2}, Pages = {258-262}, Booktitle = {Flora of North America North of Mexico}, Publisher = {Oxford University Press}, Editor = {Flora of North America Editorial Committee}, Year = {1993}, Key = {fds16676} } @article{fds230131, Author = {Pryer, KM}, Title = {The status of Gymnocarpium heterosporum and Gymnocarpium robertianum in Pennsylvania.}, Journal = {American fern journal.}, Volume = {82}, Number = {1}, Pages = {34-39}, Publisher = {JSTOR}, Year = {1992}, Month = {March}, ISSN = {0002-8444}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1992HQ04200006&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Doi = {10.2307/1547760}, Key = {fds230131} } @article{fds230132, Author = {Pryer, KM}, Title = {The limestone oak fern: new to the flora of Manitoba}, Journal = {Blue Jay}, Volume = {48}, Number = {4}, Pages = {192-195}, Publisher = {University of Alberta Libraries}, Year = {1990}, url = {http://dx.doi.org/10.29173/bluejay4791}, Doi = {10.29173/bluejay4791}, Key = {fds230132} } @article{fds353371, Author = {Argus, GW and Pryer, KM}, Title = {Les plantes vasculaires rares du Canada(notre patrimoine naturel)}, Publisher = {Musée canadien de la nature}, Year = {1990}, Key = {fds353371} } @article{fds353372, Author = {Argus, GW and Pryer, KM}, Title = {Les plantes vasculaires rares du Canada}, Journal = {Musée canadien de la nature, Ottawa, Ont}, Year = {1990}, Key = {fds353372} } @article{fds230134, Author = {Pryer, KM and Phillippe, LR}, Title = {A synopsis of the genus Sanicula (Apiaceae) in eastern Canada}, Journal = {Canadian Journal of Botany}, Volume = {67}, Number = {3}, Pages = {694-707}, Publisher = {Canadian Science Publishing}, Year = {1989}, Month = {January}, ISSN = {0008-4026}, url = {http://dx.doi.org/10.1139/b89-093}, Abstract = {Four species and 2 varieties of these native woodland umbellifers are recognized. A key to the taxa, comparative descriptions of diagnostic characters, and notes on the taxonomy, distribution, habitat, and rare status are provided. Eastern Canadian dot maps and North American range maps are included for each taxon. -from Authors}, Doi = {10.1139/b89-093}, Key = {fds230134} } @article{fds353373, Author = {Pryer, KM and Windham, MD}, Title = {A re-examination of Gymnocarpium dryopteris (L.) Newman in North America}, Journal = {Amer. J. Bot}, Volume = {75}, Pages = {142}, Year = {1988}, Key = {fds353373} } @article{fds353374, Author = {PRYER, K and ARGUS, G and HABER, E}, Title = {THE CANADIAN RARE AND ENDANGERED PLANTS PROJECT-A BAKERS DECADE LATER}, Journal = {AMERICAN JOURNAL OF BOTANY}, Volume = {73}, Number = {5}, Pages = {781-782}, Publisher = {BOTANICAL SOC AMER INC OHIO STATE UNIV-DEPT BOTANY 1735 NEIL AVE, COLUMBUS …}, Year = {1986}, Key = {fds353374} } @article{fds230135, Author = {McNeill, J and Pryer, KM}, Title = {THE STATUS AND TYPIFICATION OF PHEGOPTERIS AND GYMNOCARPIUM}, Journal = {TAXON}, Volume = {34}, Number = {1}, Pages = {136-143}, Publisher = {Wiley}, Year = {1985}, Month = {February}, ISSN = {0040-0262}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1985ADK6300024&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Abstract = {<jats:title>Summary</jats:title><jats:p>It is concluded that <jats:italic>Phegopteris</jats:italic> Fée (Thelypteridaceae) is an apparently nomenclaturally superfluous name, because, when published, it included what is now established to be the type of <jats:italic>Gymnocarpium</jats:italic> Newman. Despite this, <jats:italic>Phegopteris</jats:italic> is a legitimate name as it has a legitimate basionym; it must, however, be cited as <jats:italic>Phegopteris</jats:italic> (Presl) Fée. The protologue of the basionym of <jats:italic>Gymnocarpium dryopteris</jats:italic> does not point to any specimen that could serve as type of that name, and hence of <jats:italic>Gymnocarpium</jats:italic> Newman (Athyriaceae). Moreover, there is no eligible specimen in the Linnaean collections in London, Stockholm or Paris. There is, however, a Burser specimen at Uppsala with a Bauhin name that is not cited in any Linnaean work. This specimen was determined by Linnaeus in his catalogue of the Burser herbarium as <jats:italic>Polypodium dryopteris</jats:italic>, the basionym of <jats:italic>G. dryopteris</jats:italic>, and this is designated as the lectotype. This lectotypification preserves existing usage of both the generic and specific names.</jats:p>}, Doi = {10.2307/1221578}, Key = {fds230135} } @article{fds230136, Author = {Pryer, KM and Britton, DM and McNeill, J}, Title = {A numerical analysis of chromatographic profiles in North American taxa of the fern genus Gymnocarpium}, Journal = {Canadian Journal of Botany}, Volume = {61}, Number = {10}, Pages = {2592-2602}, Publisher = {Canadian Science Publishing}, Year = {1983}, Month = {October}, ISSN = {0008-4026}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1983RR31700012&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Abstract = {<jats:p> As part of a systematic investigation of the genus Gymnocarpium in North America, a survey of chromatographic profiles in species and hybrids of the genus was initiated. It was established through cluster analysis and ordination of the phenolic data that morphologically distinguishable taxa of Gymnocarpium can be recognized by their chromatographic profiles alone. These data provide supportive evidence for the recognition of G. robertianum and G. jessoense ssp. parvulum as distinct taxa and for the hybrid status of G. × intermedium. They also suggest that, as currently circumscribed, G. jessoense ssp. jessoense is a heterogeneous taxon. </jats:p>}, Doi = {10.1139/b83-285}, Key = {fds230136} } @article{fds230137, Author = {Pryer, KM and Britton, DM}, Title = {Spore studies in the genus Gymnocarpium}, Journal = {Canadian Journal of Botany}, Volume = {61}, Number = {2}, Pages = {377-388}, Publisher = {Canadian Science Publishing}, Year = {1983}, Month = {February}, ISSN = {0008-4026}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:A1983QF41300001&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Abstract = {<jats:p> Scanning electron microscopy was used to examine the variation in perispore characters within and between six currently recognized taxa of the genus Gymnocarpium Newm. in North America. Representative scanning electron micrographs are presented and depict the perispore features observed. Spores of those taxa studied here do not each possess a distinctive morphology providing practical species discrimination, rather they all exhibit similar patterns of variation. Spore size was also examined and shown to be correlated to ploidy level, permitting positive identification of the diploid taxon G. dryopteris ssp. disjunctum from the tetraploid G. dryopteris ssp. dryopteris. </jats:p>}, Doi = {10.1139/b83-045}, Key = {fds230137} } @article{fds353376, Author = {Argus, GW and Pryer, KM and White, DJ and Keddy, CJ}, Title = {1987}, Journal = {Atlas of the rare vascular plants of Ontario}, Volume = {4}, Year = {1982}, Key = {fds353376} } @article{fds230138, Author = {Sarvela, J and Britton, DM and Pryer, K}, Title = {Studies on the Gymnocarpium robertianum complex in North America}, Journal = {Rhodora}, Volume = {83}, Number = {835}, Pages = {421-431}, Publisher = {New England Botanical Club}, Year = {1981}, Key = {fds230138} } @article{fds230139, Author = {Chase, R and Pryer, K and Baker, R and Madison, D}, Title = {Responses to conspecific chemical stimuli in the treatment snail Achatina fulica (Pulmonata: Sigmurethra).}, Journal = {Behavioral biology}, Volume = {22}, Number = {3}, Pages = {302-315}, Year = {1978}, Month = {March}, ISSN = {0091-6773}, url = {http://dx.doi.org/10.1016/s0091-6773(78)92366-0}, Abstract = {The giant African snail, Achatina fulica, followed trails made with the mucus of A. fulica, but did not follow those consisting of mucus from Otala vermiculata. In olfactometer experiments, A. fulica and Helix aperta oriented preferentially toward the odor of their own species when both odors were presented simultaneously. Species specificity was less pronounced when the odor of O. vermiculata was paired with either of the other two snail odors. Sexually mature A. fulica that had been housed individually for 30 days prior to testing followed mucus trails better than did similar snails housed collectively. Immature A. fulica did not follow trails better after isolation, but showed a facilitative effect of isolation on conspecific orientation in the olfactometer. Three-week-old snails, maintained in individual containers from the time of hatching, also oriented preferentially toward conspecific odors. © 1978 Academic Press, Inc.}, Doi = {10.1016/s0091-6773(78)92366-0}, Key = {fds230139} } %% Papers Submitted @article{fds220952, Author = {Sigel, E.M. and M.D. Windham and C.H. Haufler and K.M. Pryer}, Title = {Phylogeny, divergence time estimates, and phylogeography for the diploid species of the Polypodium vulgare complex}, Journal = {Systematic Botany, in review}, Year = {2013}, Key = {fds220952} } @article{fds220954, Author = {Rothfels, C.J. and A.K. Johnson and P.H. Hovenkamp and D.L. Swofford and H.C. Roskam and C.R. Fraser-Jenkins and M.D. Windham and K.M. Pryer}, Title = {A natural intergeneric hybrid between parental lineages that diverged from one another over 50 million years ago}, Journal = {Evolution, in review}, Year = {2013}, Key = {fds220954} } %% Book Chapters @misc{fds219919, Author = {Wang, Z.R. and K.M. Pryer}, Title = {Gymnocarpium}, Pages = {257-259}, Booktitle = {Z.Y. Wu, P.H. Raven & D.Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press}, Year = {2013}, Key = {fds219919} } @misc{fds219920, Author = {Wang, Z.R. and C.H. Haufler and K.M. Pryer and M. Kato}, Title = {Cystopteridaceae}, Pages = {257}, Booktitle = {Z.Y. Wu, P.H. Raven & D.Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press}, Year = {2013}, Key = {fds219920} } @misc{fds71728, Author = {Pryer, K.M. and E. Schuettpelz}, Title = {Ferns}, Pages = {153-156}, Booktitle = {The Timetree of Life, Chapter 14 in S.B. Hedges and S. Kumar (eds.)}, Publisher = {Oxford University Press, New York}, Year = {2009}, Key = {fds71728} } @misc{fds230058, Author = {Schuettpelz, E and Pryer, KM}, Title = {Fern phylogeny}, Pages = {395-416}, Booktitle = {Biology and Evolution of Ferns and Lycophytes}, Publisher = {Cambridge University Press}, Year = {2008}, Month = {January}, ISBN = {9780521874113}, url = {http://dx.doi.org/10.1017/CBO9780511541827.016}, Abstract = {Introduction As a consequence of employing DNA sequence data and phylogenetic approaches, unprecedented progress has been made in recent years toward a full understanding of the fern tree of life. At the broadest level, molecular phylogenetic analyses have helped to elucidate which of the so-called “fern allies” are indeed ferns, and which are only distantly related (Nickrent et al., 2000; Pryer et al., 2001a; Wikström and Pryer, 2005; Qiu et al., 2006). Slightly more focused analyses have revealed the composition of, and relationships among, the major extant fern clades (Hasebe et al., 1995; Wolf, 1997; Pryer et al., 2004b; Schneider et al., 2004c; Schuettpelz et al., 2006; Schuettpelz and Pryer, 2007). A plethora of analyses, at an even finer scale, has uncovered some of the most detailed associations (numerous references cited below). Together, these studies have helped to answer many long-standing questions in fern systematics. In this chapter, a brief synopsis of vascular plant relationships - as currently understood - is initially provided to place ferns within a broader phylogenetic framework. This is followed by an overview of fern phylogeny, with most attention devoted to the leptosporangiate clade that accounts for the bulk of extant fern diversity. Discussion of finer scale relationships is generally avoided; instead, the reader is directed to the relevant literature, where more detailed information can be found.}, Doi = {10.1017/CBO9780511541827.016}, Key = {fds230058} } @misc{fds230059, Author = {Smith, AR and Pryer, KATHLEENM and Schuettpelz, ERIC and Korall, P and Schneider, HARALD and Wolf, PG}, Title = {Fern classification}, Pages = {417-467}, Booktitle = {Biology and Evolution of Ferns and Lycophytes}, Publisher = {Cambridge University Press}, Editor = {Ranker, T.A. and C.H. Haufler (eds.).}, Year = {2008}, ISBN = {9780521874113}, url = {http://dx.doi.org/10.1017/CBO9780511541827.017}, Abstract = {Introduction and historical summary Over the past 70 years, many fern classifications, nearly all based on morphology, most explicitly or implicitly phylogenetic, have been proposed. The most complete and commonly used classifications, some intended primarily as herbarium (filing) schemes, are summarized in Table 16.1, and include: Christensen (1938), Copeland (1947), Holttum (1947, 1949), Nayar (1970), Bierhorst (1971), Crabbe et al. (1975), Pichi Sermolli (1977), Ching (1978), Tryon and Tryon (1982), Kramer (in Kubitzki, 1990), Hennipman (1996), and Stevenson and Loconte (1996). Other classifications or trees implying relationships, some with a regional focus, include Bower (1926), Ching (1940), Dickason (1946), Wagner (1969), Tagawa and Iwatsuki (1972), Holttum (1973), and M.ckel (1974). Tryon (1952) and Pichi Sermolli (1973) reviewed and reproduced many of these and still earlier classifications, and Pichi Sermolli (1970, 1981, 1982, 1986) also summarized information on family names of ferns. Smith (1996) provided a summary and discussion of recent classifications. With the advent of cladistic methods and molecular sequencing techniques, there has been an increased interest in classifications reflecting evolutionary relationships. Phylogenetic studies robustly support a basal dichotomy within vascular plants, separating the lycophytes (less than 1% of extant vascular plants) from the euphyllophytes (Figure 16.1; Raubeson and Jansen, 1992, Kenrick and Crane, 1997; Pryer et al., 2001a, 2004a, 2004b; Qiu et al., 2006). Living euphyllophytes, in turn, comprise two major clades: spermatophytes (seed plants), which are in excess of 260000 species (Thorne, 2002; Scotland and Wortley, 2003), and ferns (sensu Pryer et al. 2004b), with about 9000 species, including horsetails, whisk ferns, and all eusporangiate and leptosporangiate ferns.}, Doi = {10.1017/CBO9780511541827.017}, Key = {fds230059} } @misc{fds230075, Author = {Pryer, KM and Schneider, H and Magallón, S}, Title = {The radiation of vascular plants}, Pages = {138-153}, Booktitle = {Assembling the Tree of Life}, Publisher = {Oxford University Press New York}, Editor = {Cracraft, J and Donoghue, MJ}, Year = {2004}, ISBN = {0-19-517234-5}, url = {http://gateway.webofknowledge.com/gateway/Gateway.cgi?GWVersion=2&SrcApp=PARTNER_APP&SrcAuth=LinksAMR&KeyUT=WOS:000225766900011&DestLinkType=FullRecord&DestApp=ALL_WOS&UsrCustomerID=47d3190e77e5a3a53558812f597b0b92}, Key = {fds230075} } @misc{fds353359, Author = {Schneider, H and Pryer, KM and Cranfill, R and Smith, AR and Wolf, PG}, Title = {Evolution of vascular plant body plans: a phylogenetic perspective}, Pages = {330-364}, Booktitle = {Developmental Genetics and Plant Evolution}, Publisher = {Taylor and Francis London}, Editor = {Q.C.B. Cronk and R.M. Bateman and J.A. Harris}, Year = {2002}, Key = {fds353359} } @misc{fds353311, Author = {Wolf, PG and Pryer, KM and Smith, AR and Hasebe, M}, Title = {Phylogenetic studies of extant pteridophytes}, Pages = {541-556}, Booktitle = {Molecular systematics of plants II}, Publisher = {Springer, Boston, MA}, Editor = {D.E. Soltis and P.S. Soltis and J. J. Doyle}, Year = {1998}, ISBN = {9780412111310}, url = {http://dx.doi.org/10.1007/978-1-4615-5419-6_19}, Doi = {10.1007/978-1-4615-5419-6_19}, Key = {fds353311} } %% Published Abstracts @misc{fds219932, Author = {Li, F.-W. and C.J. Rothfels and A. Larsson and E.M. Sigel and L. Huiet and P. Korall and M. Ruhsam and D. Stevenson and S. Graham and G.K.-S. Wong and K.M. Pryer}, Title = {Mining fern transcriptome data for low-copy nuclear markers}, Year = {2013}, Abstract = {http://2013.botanyconference.org/engine/search/index.php?func=detail&aid=626}, Key = {fds219932} } @misc{fds219933, Author = {Johnson, A.K. and A.L. Grusz and J.B. Beck and K.M. Pryer and M.D. Windham}, Title = {So, if they are evolutionary dead-ends, why are apomictic cheilanthoid ferns more widely distributed than their sexual diploid progenitors?}, Year = {2013}, Abstract = {http://2013.botanyconference.org/engine/search/index.php?func=detail&aid=803}, Key = {fds219933} } @misc{fds219934, Author = {Sigel, E.M. and J. Der and C.W. De Pamphilis and K.M. Pryer}, Title = {Time after time: Comparing homeolog expression patterns between two independent origins of the allotetraploid fern Polypodium hesperium.}, Year = {2013}, Abstract = {http://2013.botanyconference.org/engine/search/index.php?func=detail&aid=651}, Key = {fds219934} } @misc{fds212621, Author = {Grusz, A.L. and M.D. Windham and K.M. Pryer}, Title = {Using next-generation sequencing to develop microsatellite markers in ferns}, Year = {2012}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=946}, Key = {fds212621} } @misc{fds212623, Author = {Beck, J. and J. Allison and K.M. Pryer and M.D. Windham}, Title = {Identifying multiple origins of polyploid taxa: a multilocus study of the hybrid cloak fern (Astrolepis integerrima; Pteridaceae).}, Year = {2012}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=293}, Key = {fds212623} } @misc{fds212624, Author = {Sigel, E.M. and M.D. Windham and C.H. Haufler and K.M. Pryer}, Title = {Reassessing phylogenetic relationships in the Polypodium vulgare complex (Polypodiaceae) using nuclear gapCp sequence data}, Year = {2012}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=890}, Key = {fds212624} } @misc{fds212625, Author = {Huiet, R.L. and K.M. Pryer and C.H. Haufler and M.D. Windham}, Title = {Simplifying the complex: an integrative analysis using cytogenetics, morphology, and DNA sequence data to resolve relationships within the Pellaea wrightiana complex (Pteridaceae).}, Year = {2012}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=651}, Key = {fds212625} } @misc{fds197476, Author = {Li, F.-W. and L.-Y. Kuo and C.J. Rothfels and A. Ebihara and W.-L. Chiou, M.D. Windham and K.M. Pryer}, Title = {The global two-locus plant barcode: a first evaluation of its utility across ferns.}, Year = {2011}, url = {http://www.ibc2011.com/downloads/IBC2011_Abstract_Book.pdf}, Key = {fds197476} } @misc{fds197468, Author = {Johnson, A.K.* and C.J. Rothfels and A.L. Grusz and E.M. Sigel and M.D. Windham and K.M. Pryer}, Title = {Sporophytes and gametophytes of notholaenid ferns (Pteridaceae) show correlated presence/absence of farina.}, Year = {2011}, url = {http://2011.botanyconference.org/engine/search/index.php?func=detail&aid=536}, Key = {fds197468} } @misc{fds197469, Author = {Hooper, E. and G. Yatskievych and L. Huiet and M.D. Windham and K.M. Pryer}, Title = {Into or out of Africa? What do molecular data reveal about the identity and biogeographic origin of Aleuritopteris farinosa (Forssk.) Fee (Pteridaceae)?}, Year = {2011}, url = {http://2011.botanyconference.org/engine/search/index.php?func=detail&aid=723}, Key = {fds197469} } @misc{fds197470, Author = {Li, F.-W. and L.-Y. Kuo and C.J. Rothfels and A. Ebihara and W.-L. Chiou, M.D. Windham and K.M. Pryer}, Title = {rbcL and matK earn a thumbs up as the core DNA barcode for ferns}, Year = {2011}, url = {http://2011.botanyconference.org/engine/search/index.php?func=detail&aid=669}, Key = {fds197470} } @misc{fds184015, Author = {Grusz, AL, MD Windham and KM Pryer}, Title = {Examining the role of apomixis in the evolution of desert-adapted ferns. Invited symposium presentation.}, Year = {2010}, Abstract = {http://2010.botanyconference.org/engine/search/index.php?func=detail&aid=717}, Key = {fds184015} } @misc{fds184017, Author = {Nagalingum, N and R Lupia and KM Pryer}, Title = {The importance of paleobotanical and herbarium collections in understanding the evolution of heterosporous ferns. Invited symposium presentation.}, Year = {2010}, Abstract = {http://2010.botanyconference.org/engine/search/index.php?func=detail&aid=564}, Key = {fds184017} } @misc{fds184018, Author = {Sigel, EM, MD Windham and KM Pryer}, Title = {Using spore data to infer ploidy and reproductive mode in land plants. Invited symposium presentation.}, Year = {2010}, Abstract = {http://2010.botanyconference.org/engine/search/683.html}, Key = {fds184018} } @misc{fds184019, Author = {Rothfels, CJ and E Schuettpelz and KM Pryer}, Title = {An accelerated rate of molecular evolution spans all three genomes in vittarioid ferns.}, Year = {2010}, Abstract = {http://2010.evolutionmeeting.org/search/index.php?func=detail&aid=771}, Key = {fds184019} } @misc{fds184020, Author = {Schuettpelz, E, MD Windham and KM Pryer}, Title = {Estimating divergence times and ancestral centers of diversity for a xeric-adapted fern clade.}, Year = {2010}, Abstract = {http://2010.evolutionmeeting.org/search/index.php?func=detail&aid=652}, Key = {fds184020} } @misc{fds168402, Author = {Grusz, A. L. and M.D. Windham and K.M. Pryer}, Title = {A Cheilanthes by any other name: Evolutionary complexity in the New World myriopterid clade (Pteridaceae).}, Year = {2009}, Abstract = {http://2009.botanyconference.org/engine/search/index.php?func=detail&aid=892}, Key = {fds168402} } @misc{fds168403, Author = {Sigel, E. M. and M.D. Windham and K.M. Pryer}, Title = {To have or have not: Using farina to delineate major clades within the false cloak ferns (Argyrochosma).}, Year = {2009}, Abstract = {http://2009.botanyconference.org/engine/search/index.php?func=detail&aid=712}, Key = {fds168403} } @misc{fds168404, Author = {Rothfels, C.J. and M.D. Windham and K.M. Pryer}, Title = {New insights into the relationships of Cystopteris, Acystopteris, and Gymnocarpium.}, Year = {2009}, Abstract = {http://2009.botanyconference.org/engine/search/index.php?func=detail&aid=625}, Key = {fds168404} } @misc{fds168405, Author = {Schuettpelz, E. and K.M. Pryer and M.D. Windham}, Title = {A phylogenetic approach to species delimitation in the fern genus Pentagramma (Pteridaceae).}, Year = {2009}, Abstract = {http://2009.botanyconference.org/engine/search/index.php?func=detail&aid=511}, Key = {fds168405} } @misc{fds168406, Author = {Beck, J. and M.D. Windham and K.M. Pryer}, Title = {Tempo and mode of evolution in the star-scaled cloak ferns (Astrolepis).}, Year = {2009}, Abstract = {http://2009.botanyconference.org/engine/search/index.php?func=detail&aid=166}, Key = {fds168406} } @misc{fds152862, Author = {Beck, J. and M.D. Windham and K.M. Pryer}, Title = {Investigating the early stages of polyploid evolution in the star-scaled cloak ferns (Astrolepis)}, Year = {2008}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=105}, Key = {fds152862} } @misc{fds152863, Author = {Windham, M.D. and J. Beck and A.L. Grusz and L. Huiet and C. Rothfels and E. Schuettpelz and G. Yatskievych and K.M. Pryer}, Title = {Using plastid and nuclear DNA sequences to redraw generic boundaries and demystify species complexes in cheilanthoid ferns}, Year = {2008}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=686}, Key = {fds152863} } @misc{fds152864, Author = {Pryer, K.M. and E.Y. Butler and D.R. Farrar and R. Moran and J.J. Schneller, E. Schuettpelz and J.E. Watkins, Jr. and M.D. Windham}, Title = {On the importance of portraying the plant life cycle accurately: ferns as a case study}, Year = {2008}, Abstract = {http://www.botanyconference.org/engine/search/index.php?func=detail&aid=848}, Key = {fds152864} } @misc{fds152848, Author = {Schuettpelz E. and H. Schneider and L. Huiet and M.D. Windham and K.M. Pryer.}, Title = {A molecular phylogeny of pteroid ferns: rampant paraphyly and polyphyly revealed.}, Year = {2007}, Abstract = {http://www.2007.botanyconference.org/engine/search/index.php?func=detail&aid=1604}, Key = {fds152848} } @misc{fds152847, Author = {Rothfels, C.J. and M.D. Windham and K.M. Pryer and J.S. Metzgar and A.L. Grusz.}, Title = {Making Fronds in the Desert: Phylogenetics of Farinose Ferns (Notholaena: Pteridaceae).}, Year = {2007}, Abstract = {http://www.2007.botanyconference.org/engine/search/index.php?func=detail&aid=1922}, Key = {fds152847} } @misc{fds152846, Author = {Pryer, K.M. and E. Schuettpelz}, Title = {Ancient Origins and recent radiations in the evolutionary history of ferns. Invited symposium presentation: Deep Time: Integrating Paleobotany and Phylogenetics.}, Year = {2007}, Abstract = {http://www.2007.botanyconference.org/engine/search/index.php?func=detail&aid=2184}, Key = {fds152846} } @misc{fds152849, Author = {Kleist, A.C. and C.L. Nelson and J.M.O. Geiger and P. Korall and T.A. Ranker and K.M. Pryer}, Title = {Alternate pathways of fern dispersal to the Hawaiian Islands, Part 3: Cibotium}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=579}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=579}, Key = {fds152849} } @misc{fds152850, Author = {Schneider, H. and K.M. Pryer and A.R. Smith.}, Title = {Green spore evolution in ferns}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=141}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=141}, Key = {fds152850} } @misc{fds152851, Author = {Metzgar, J.S. and J.E. Skog and E. Zimmer and K.M. Pryer}, Title = {Is Osmunda paraphyletic?}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=501}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=501}, Key = {fds152851} } @misc{fds152852, Author = {Lupia, R. and H. Schneider and N.S. Nagalingum and K.M. Pryer}, Title = {Jurassic origin for the Salviniaceae: the last word or just the first?}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=535}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=535}, Key = {fds152852} } @misc{fds152853, Author = {Korall, P. and J.S. Metzgar and D.S. Conant and H. Schneider and K.M. Pryer}, Title = {Phylogeny of scaly tree ferns (Cyatheaceae)}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=289}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=289}, Key = {fds152853} } @misc{fds152854, Author = {Grusz, A.L. and M.D. Windham and J.S. Metzgar and K.M. Pryer}, Title = {Polyploids and reticulate voids: the Cheilanthes fendleri complex revisited}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=384}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=384}, Key = {fds152854} } @misc{fds152855, Author = {Nagalingum, N.S. and K.M. Pryer and H. Schneider and D. Hearn}, Title = {The age of heterosporous ferns as estimated by molecular divergence dating and the fossil record}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=520}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=520}, Key = {fds152855} } @misc{fds152856, Author = {K.M. Pryer and Schuettpelz, E.}, Title = {Toward a comprehensive phylogeny of extant ferns}, Year = {2006}, url = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=469}, Abstract = {http://www.2006.botanyconference.org/engine/search/index.php?func=detail&aid=469}, Key = {fds152856} } @misc{fds152857, Author = {N. Nagalingum and K.M. Pryer and H. Schneider and M.D. Nowak and D. Hearn, R. Lupia}, Title = {Dating the Marsileaceae: evolutionary and biogeographical implications. Symposium 12.7.2 on DATING DIVERGENCE OF ANGOIOSPERM RADIATIONS: PROGRESS AND PROSPECTS.}, Year = {2005}, Month = {Summer}, url = {http://www.ibc2005.ac.at/program/final_program_IBC2005.pdf}, Key = {fds152857} } @misc{fds152858, Author = {Pryer, K.M. and E. Schuettpelz}, Title = {Evolutionary history of ferns: ancient origins and recent radiations. Symposium 9.5.5 on EARLY EVOLUTION: MOLECULAR PHYLOGENETICS AND ORGANELLAR GENOMICS.}, Year = {2005}, Month = {Summer}, url = {http://www.ibc2005.ac.at/program/final_program_IBC2005.pdf}, Key = {fds152858} } @misc{fds152859, Author = {Hennequin, S. and E. Schuettpelz and K.M. Pryer and J.-Y. Dubuisson}, Title = {Fast and slow filmy ferns: molecular rate heterogeneity a chloroplast-wide phenomenon in the Hymenophyllaceae. Symposium 6.9.3 on DIVERSIFICATION OF SEED-FREE VASCULAR PLANTS IN THE SHADOW OF ANGIOSPERMS}, Year = {2005}, Month = {Summer}, url = {http://www.ibc2005.ac.at/program/final_program_IBC2005.pdf}, Key = {fds152859} } @misc{fds152860, Author = {Nagalingum, N.S. and H. Schneider and K.M. Pryer}, Title = {Comparative morphology of reproductive structures in heterosporous water ferns: the homology of the sporocarp reexamined}, Year = {2004}, Abstract = {http://www.botanyconference.org/engine/search/ index.php?func=detail&aid=634}, Key = {fds152860} } @misc{fds152861, Author = {Schuettpelz, E. and H. Schneider and K.M. Pryer}, Title = {An evolutionary time-scale for ferns: ancient origins and recent radiations}, Year = {2004}, Abstract = {http://www.botanyconference.org/engine/search/ index.php?func=detail&aid=249}, Key = {fds152861} } %% Other @misc{fds212622, Author = {Chai, N. and T. Shao and R.L. Huiet and M. Kim and C.J. Rothfels and J. Golan and K.M. Pryer}, Title = {DUKE Vascular Type Specimens - Digital Images & Protologues}, Year = {2012}, url = {http://herbarium.duke.edu/collections/vasculars/typespecimens}, Key = {fds212622} } @misc{fds44577, Author = {Pryer, K.M.}, Title = {Fern DNA Database – an online database of vouchered DNA sequence data for ferns}, Year = {2010}, url = {http://www.biology.duke.edu/pryerlab/ferndb/}, Key = {fds44577} } @misc{fds184615, Author = {Nagalingum, N. and E. Schuettpelz and K.M. Pryer}, Title = {Toward a comprehensive fern tree of life: exploration and training in the forests of Malaysia}, Year = {2009}, Abstract = {http://malaysianferns.pryerlab.net/}, Key = {fds184615} } @misc{fds170472, Author = {Pryer, K.M. and A.R. Smith and C.J. Rothfels}, Title = {Polypodiopsida Cronquist, Takht. & Zimmerm. 1966. Ferns. In Maddison, D.R. and W. P. Maddison. The Tree of Life: A distributed Internet project containing information about phylogeny and biodiversity.}, Year = {2009}, url = {http://tolweb.org/Polypodiidae/21666}, Abstract = {http://tolweb.org/tree?group=Polypodiopsida&contgroup=Embryophytes}, Key = {fds170472} } @misc{fds36198, Author = {K.M. Pryer}, Title = {Fern Image Database}, Year = {2007}, url = {http://www.pryerlab.net/img_dbase/index.shtml}, Key = {fds36198} } @misc{fds44573, Author = {Windham, M.D. and D. Hearn and J. Metzgar and E. Schuettpelz and K.M. Pryer}, Title = {Ferns of Arizona (an online flora)}, Year = {2005}, Month = {Summer}, url = {http://AZferns.org}, Key = {fds44573} } @misc{fds16763, Author = {Schneider, H. and K.M. Pryer}, Title = {Convergent evolution in vegetative and reproductive characters of aquatic vascular plants}, Year = {2003}, Key = {fds16763} } @misc{fds16765, Author = {Schneider, H. and R. Cranfill and E.J. Schuettpelz and S. Magallón and K.M. Pryer}, Title = {Derived ferns diversified in the shadow of angiosperms: evidence from the fossil record, phylogenetic patterns, and divergence time estimates}, Year = {2003}, Key = {fds16765} } @misc{fds16767, Author = {Chikarmane, S. and T. Rehse and K.M. Pryer}, Title = {Tracing the cultural and botanical origins of turmeric (Curcuma longa L.)}, Year = {2003}, Key = {fds16767} } @misc{fds16768, Author = {Schuettpelz, E.J. and K.M. Pryer}, Title = {Characterizing molecular rate heterogeneity between two major lineages of filmy ferns (Hymenophyllaceae)}, Year = {2003}, Key = {fds16768} } @misc{fds44578, Author = {K.M. Pryer}, Title = {Fern Image Database}, Year = {2003}, url = {http://www.pryerlab.net/img_dbase/index.shtml}, Key = {fds44578} } @misc{fds16775, Title = {Atlas of the rare vascular plants of Ontario. Part 4.}, Editor = {Pryer, K.M. and G.W. Argus}, Year = {2003}, Key = {fds16775} } @misc{fds16761, Author = {Pryer, K.M. and H. Schneider}, Title = {A phylogeny for extant heterosporous ferns}, Year = {2002}, Key = {fds16761} } @misc{fds16762, Author = {Schuettpelz, E. and H. Schneider and K. M. Pryer}, Title = {The phylogenetic history of Pterozonium (Pteridaceae) revisited: from groundplan divergence to maximum parsimony}, Year = {2002}, Key = {fds16762} } @misc{fds16759, Author = {Schneider, H. and K. M. Pryer and R. Lupia}, Title = {A comparative analysis of structure and function of spores in extant heterosporous ferns (Salviniales)}, Year = {2001}, Key = {fds16759} } @misc{fds16760, Author = {Des Marais and D.L., K.M. Pryer and A.R. Smith}, Title = {Phylogeny, character evolution, and biogeography of extant horsetails (Equisetum)}, Year = {2001}, Key = {fds16760} } @misc{fds16758, Author = {Schneider, H. and K.M. Pryer}, Title = {Spore morphology of heterosporous ferns and its possible implications for understanding the evolution of the seed habit}, Journal = {American Journal of Botany}, Volume = {87 (Suppl)}, Pages = {31-32}, Year = {2000}, Key = {fds16758} } @misc{fds16748, Author = {Pryer, K.M. and J.-Y. DuBuisson}, Title = {Symposium: Fern phylogeny with an emphasis on relationships of basal lineages}, Journal = {XVI International Botanical Congress Abstracts}, Pages = {91}, Year = {1999}, Key = {fds16748} } @misc{fds16749, Author = {DuBuisson, J.-Y. and J.S. Hunt and K.M. Pryer and A.R. Smith}, Title = {Phylogeny of filmy ferns (Hymenophyllaceae)}, Journal = {XVI International Botanical Congress Abstracts}, Pages = {91}, Year = {1999}, Key = {fds16749} } @misc{fds16752, Author = {Hunt, J.S. and K.M. Pryer and A. Vaghani and A.R. Smith and P.G. Wolf}, Title = {"Fern DNA Database": using FileMaker Pro to coordinate DNA-availability, DNA-sequence data, and voucher and source information for large-scale and collaborative phylogenetic studies}, Journal = {XVI International Botanical Congress Abstracts}, Pages = {720}, Year = {1999}, Key = {fds16752} } @misc{fds16753, Author = {Moeser, G.M. and R. Lupia and H. Schneider and K.M. Pryer}, Title = {Marsileaceae sporocarps and spores from the Late Cretaceous}, Journal = {XVI International Botanical Congress Abstracts}, Pages = {338}, Year = {1999}, Key = {fds16753} } @misc{fds16755, Author = {Schneider, H. and K.M.Pryer, J.S. Hunt and L. Sappelsa and P.G. Wolf and S. D. Sipes and A.R. Smith}, Title = {Basal tracheophytes and the phylogeny of “pteridophyte” lineages – inferred from four large data sets: morphology, 18S nrDNA, rbcL, and atpB}, Journal = {14th Symposium for Biodiversität und Evolutionsbiologie, Jena, Germany}, Pages = {163}, Year = {1999}, Key = {fds16755} } @misc{fds16756, Author = {Schneider, H. and K.M. Pryer and J.S. Hunt and L. Sappelsa and P.G. Wolf and S. D. Sipes and A.R. Smith}, Title = {Basal tracheophytes and the phylogeny of “pteridophyte” lineages – inferred from four large data sets: morphology, 18S nrDNA, rbcL, and atpB, Göttingen, Germany}, Journal = {XVIIIth meeting of the Willi Hennig Society}, Pages = {46}, Year = {1999}, Key = {fds16756} } @misc{fds16747, Author = {Palmer, J.D. and S. Turner and K.M. Pryer}, Title = {Cyanobacterial origin of chloroplasts: once or more than once?}, Journal = {Journal of Eukaryotic Microbiology}, Volume = {46}, Pages = {2A}, Year = {1998}, Key = {fds16747} } @misc{fds16744, Author = {Pryer, K.M. and A.R. Smith and G. Rothwell and P. Kenrick}, Title = {Symposium: Relationships and fossil history of ferns}, Journal = {American Journal of Botany}, Volume = {84 (Suppl)}, Pages = {157-158}, Year = {1997}, Key = {fds16744} } @misc{fds16746, Author = {Pryer, K.M. and A.R. Smith}, Title = {Character evolution and fern phylogeny}, Journal = {American Journal of Botany}, Volume = {84 (Suppl)}, Pages = {159}, Year = {1997}, Key = {fds16746} } @misc{fds16743, Author = {Pryer, K.M.}, Title = {Ontogeny and phylogeny of marsileaceous ferns: evidence for heterochronic evolution}, Journal = {American Journal of Botany}, Volume = {83 (Suppl)}, Pages = {130}, Year = {1996}, Key = {fds16743} } @misc{fds16738, Author = {Hasebe, M. and P.G. Wolf and W. Hauk and J.R. Manhart and C.H. Haufler and R. Sano and G.J. Gastony and N. Crane and K.M. Pryer and N. Murakami and J. Yokoyama and M. Ito}, Title = {A global analysis of fern phylogeny based on rbcL nucleotide sequences}, Journal = {American Journal of Botany}, Volume = {81 (Suppl)}, Pages = {120-121}, Year = {1994}, Key = {fds16738} } @misc{fds16741, Author = {Pryer, K.M. and F.M. Lutzoni and A.R. Smith}, Title = {Towards a fern phylogeny: integrating morphology and molecules}, Journal = {American Journal of Botany}, Volume = {81 (Suppl)}, Pages = {122}, Year = {1994}, Key = {fds16741} } @misc{fds16737, Author = {Pryer, K.M}, Title = {Ontogeny and phylogeny in the aquatic fern family Marsileaceae: cladistic analysis of morphological and molecular data}, Journal = {American Journal of Botany}, Volume = {80 (Suppl)}, Pages = {172}, Year = {1993}, Key = {fds16737} } @misc{fds16736, Author = {Pryer, K.M. and M.D. Windham}, Title = {A re-examination of Gymnocarpium dryopteris (L.)}, Journal = {Newman in North America. American Journal of Botany}, Volume = {75 (Suppl)}, Pages = {142}, Year = {1988}, Key = {fds16736} } @misc{fds16735, Author = {Pryer, K.M. and G.W. Argus and E. Haber}, Title = {The Canadian Rare and Endangered Plant's Project}, Journal = {Botany '86 - Program of the 22nd Annual Meeting of the Canadian Botanical Association}, Pages = {34}, Year = {1986}, Key = {fds16735} } @misc{fds16731, Author = {Pryer, K.M. and D.M. Britton and J. McNeill}, Title = {Hybridization in the fern genus Gymnocarpium Newman in North America}, Journal = {American Journal of Botany}, Volume = {71(Suppl)}, Pages = {142}, Year = {1984}, Key = {fds16731} } @misc{fds16727, Author = {Pryer, K.M. and D.M. Britton and J. McNeill}, Title = {Systematic studies in the genus Gymnocarpium Newman in North America}, Journal = {American Journal of Botany}, Volume = {70 (Suppl)}, Pages = {60}, Year = {1983}, Key = {fds16727} } | |
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