Refereed Publications



Schuettpelz, E.  M. D. Windham, J. Prado, E. A. Hooper, G. Yatskievych, L. Huiet, K. T. Picard and K. M. Pryer. 2024. A phylogenetically-informed generic reclassification of the hemionitid ferns (Pteridaceae: Cheilanthoideae). Taxon: in review


Pryer, K. M. 2023. Remembering George Argus: a phenomenal mentor. Canadian Botanical Association Bulletin 56: 16-17. Reprint PDF

Windham, M.D., K.T. Picard,  and K.M. Pryer. 2023. An in-depth investigation of cryptic taxonomic diversity in the rare endemic mustard Draba maguirei. American Journal of Botany. e16138. Reprint PDF

Hay, N.M., J.V. Akinwuntan, V. Cai, M.D. Windham, and K.M. Pryer. 2023. Exploring past and future distributions of the rare Appalachian oak fern using MaxEnt modeling. American Fern Journal 113: 109-125. Reprint PDF

Hay*, N.M., M.D. Windham*, T. Mandáková, M.A. Lysák, K.P. Hendriks, K. Mummenhoff, F. Lens, K.M. Pryer, and C.D. Bailey. 2023. A Hyb-Seq phylogeny of Boechera and related genera using a combination of Angiosperms353 and Brassicaceae-specific bait sets. American Journal of Botany.  *Equal contributors. Reprint PDF

Pryer, K. M. 2023. A tribute to Nathalie Seenamah Nagalingum (1975–2022).  American Fern Journal 113: 265-274. Reprint PDF.


Windham, M. D., L. Huiet, J. S. Metzgar, T. A. Ranker, G. Yatskievych, C. H. Haufler, and K. M. Pryer. 2022. Once more unto the breach, dear friends: resolving the origins and relationships of the Pellaea wrightiana hybrid complex. American Journal of Botany 109: 821–850. Reprint PDF

Windham, M.D.,  K.M. Pryer, and L. Allphin. 2022. An overview of New Mexican Boechera (Brassicaceae) including three new species and thirteen new nomenclatural combinations. Journal of the Botanical Research Institute of Texas 16: 9-24. Reprint PDF

Windham, M.D.  and K.M. Pryer. 2022. First report of Utah fragile fern (Cystopteris; Cystopteridaceae) from New Mexico, and new nomenclatural combinations in Cystopteris tenneseensis. Journal of the Botanical Research Institute of Texas 16: 1-7. Reprint PDF

Windham, M.D., K.T. Picard,  and K.M. Pryer. 2022. Myriopteris grusziae: A new species from Texas and Oklahoma segregated from the Chihuahuan Desert taxon M. scabra (Pteridaceae). Systematic Botany 47: 876-886. Reprint PDF


Grusz, A.L. , M.D. Windham, K.T. Picard, K.M. Pryer, E. Schuettpelz, and C.H. Haufler. 2021. A drought-driven model for the evolution of obligate apomixis in ferns: Evidence from pellaeids (Pteridaceae). American Journal of Botany 108: 263–283. doi:10.1002/ajb2.1611 [Invited paper for special issue on “Life Without Water ”] Reprint PDF

Fauskee, B., E.M. Sigel, K.M. Pryer, and A.L. Grusz. 2021. Variation in frequency of plastid RNA editing within Adiantum implies rapid evolution in fern plastomes. American Journal of Botany 108:  820-827.  Reprint PDF

Sosa, K., K.M. Pryer, L. Huiet, G. Yatskievych, and M.D. Windham. 2021. Cheilanthes ecuadorensis: A new species of Cheilanthes s.s. (Pteridaceae) from northern South America. Systematic Botany 46: 249-259. Reprint PDF

Heron*, K., M.D. Windham*, D.R. Farrar, and K.M. Pryer. 2021. Looking back on 130 years of fern and lycophyte research in Glacier National Park, Montana: A modern taxonomic account. American Fern Journal 111: 223-250.  *Equal contributors. Reprint PDF


Windham, M.D., K.M. Pryer, D.B. Poindexter,  F.-W. Li, C.J. Rothfels, and J.B. Beck. 2020. A step-by-step protocol for meiotic chromosome counts in flowering plants: A powerful and economical technique revisited. Applications in Plant Sciences 8(4): e11342. Reprint PDF

Kao, T.-T., C.J. Rothfels, A. Melgoza-Castillo, K.M. Pryer, and M.D. Windham. 2020. Infraspecific diversification of the star cloak fern (Notholaena standleyi) in the deserts of the United States and Mexico. American Journal of  Botany 107: 658–675. Reprint PDF

Meineke E.K., C. Tomasi, S. Yuan, and K.M. Pryer. 2020. Applying machine learning to investigate long term insect-plant interactions preserved on digitized herbarium specimens. Applications in Plant Sciences 8(6): e11369. Reprint PDF

Pryer, K.M., C. Tomasi, X. Wang, E.K. Meineke, and M.D. Windham. 2020. Using computer vision on herbarium specimen images to discriminate among closely related horsetails (Equisetum). Applications in Plant Sciences 8(6): e11372. Reprint PDF

Eily, A.N, K.M. Pryer, and F.-W. Li.  2019. A first glimpse at genes important to the Azolla–Nostoc symbiosis. Symbiosis 77: Reprint PDF

George, L.O.,  K.M. Pryer, T.-T., Kao, L. Huiet, and M.D. Windham. 2019. Baja: A new monospecific genus segregated from Cheilanthes s.l. (Pteridaceae).  Systematic Botany 44471-482. Reprint PDF

Kao, T.-T., K.M. Pryer, F.D. Freund, M.D. Windham, and C.J. Rothfels. 2019. Low-copy nuclear sequence data confirms homoplasy in farina expression in notholaenid ferns (Pteridaceae). Molecular Phylogenetics and Evolution 138: 139 -155. Reprint PDF

Yang, E.J., C.Y. Yoo, J, Liu, H. Wang, J. Cao, F.-W. Li, K.M. Pryer, T.-P. Sun, D. Weiglel, P. Zhou, M. Chen. 2019. NCP activates chloroplast transcription by controlling phytochrome-dependent dual nuclear and plastidial switches. Nature Communications 10:  2630.  Reprint PDF.

Sigel, E.M., J.P. Der, M.D. Windham, and K.M. Pryer. 2019. Expression level dominance and homeolog expression bias in recurrent origins of the allopolyploid fern Polypodium hesperium. American Fern Journal 109:  224-247. Reprint PDF

Dijkhuizen, L., P.  Brouwer, H. Bolhuis, G.-J. Reichart, N. Koppers, B. Huettel, A. Bolger, F.-W.  Li, S. Cheng, X. Liu, G.K.-S. Wong, K.M. Pryer, A. Weber, A. Bräutigam, and H. Schluepmann. 2018. Is there foul play in the leaf pocket? The metagenome of floating fern Azolla reveals endophytes that do not fix N2 but may denitrify. New Phytologist 217: 453–466. Reprint PDF

Song, M.J.B., L.-Y. Kuo, L. Huiet, K.M. Pryer, C.J. Rothfels, and F.-W. Li.  2018. A novel chloroplast gene reported for flagellate plants.  American Journal of Botany 105: 117-121. Reprint PDF   Featured inAJB News and Views 

Schuettpelz, E., G. Rouhan, K.M. Pryer, C.J. Rothfels, J. Prado, M.A. Sundue, M.D. Windham, R.C. Moran and A.R. Smith. 2018. Are there too many fern genera? Taxon 47: 473-480. Reprint PDF

Li, F.-W., P. Brouwer, L. Carretero-Paulet, S. Cheng, J. de Vries, P.-M. Delaux, A.N. Eily, N. Koppers, L.-Y. Kuo, Z. Li, M. Simenc, I. Small, E. Wafula, S. Angarita, M.S. Barker, A. Braeutigam, C. dePamphilis, S. Gould, P.S. Hosmani, Y.-M. Huang, B. Huettel, Y. Kato, X. Liu, S. Maere, R. McDowell, L.A. Mueller, K.G.J. Nierop, S.A. Rensing, T. Robison, C.J. Rothfels, E.M. Sigel, Y. Song, P.R. Timilsina, Y. Van de Peer, H. Wang, P. K.I. Wilhelmsson, P.G. Wolf, X. Xu, J.P. Der, H. Schluepmann, G.K.-S. Wong, and K.M. Pryer.  2018. Fern genomes elucidate land plant evolution and cyanobacterial symbioses. Nature Plants 4: 460–472.  Reprint PDF   COVER!

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Huiet, L., F.-W. Li, T.-T. Kao, J. Prado, A.R. Smith, E. Schuettpelz and K.M. Pryer. 2018. A worldwide phylogeny of Adiantum (Pteridaceae) reveals remarkable convergent evolution in leaf blade architecture. Taxon 67: 488–502. Reprint PDF

Sigel, E.M., E. Schuettpelz, K.M. Pryer, and J.P. Der. 2018. Overlapping patterns of gene expression between gametophyte and sporophyte phases in the fern Polypodium amorphum (Polypodiales).  Frontiers in Plant Science 9: 1450. doi: 10.3389/fpls.2018.01450. Reprint PDF

Rothfels, C.J., K.M. Pryer, and F.-W. Li. 2017. Next-generation polyploid phylogenetics: Rapid resolution of hybrid polyploid complexes using PacBio single-molecule sequencing. New Phytologist 213: 413-429Reprint PDF

Hirai, R.Y., E. Schuettpelz, L. Huiet, K.M. Pryer, A.R. Smith, and J. Prado. 2016. Phylogeny and relationships of the neotropical Adiantum raddianum group (Pteridaceae). Taxon 65: 1225–1235. Reprint PDF

Li, F.-W., L.-Y. Kuo, K.M. Pryer, and C.J. Rothfels. 2016. Genes translocated into the plastid inverted repeat show decelerated substitution rates and elevated GC content. Genome Biology and Evolution 8: 2452–2458. Reprint PDF   COVER!

Haufler, C.H., K.M. Pryer, E. Schuettpelz, E.B. Sessa, D.R. Farrar, R. Moran, J.J. Schneller, J.E. Watkins, Jr., and M.D. Windham. 2016. Sex and the single gametophyte: Revising the homosporous vascular plant life cycle in light of contemporary research. BioScience 66: 928-937. Reprint PDF

Schuettpelz, E., C.-W. Chen, M. Kessler, J.B. Pinson, G. Johnson, A. Davila, A.T. Cochran, L. Huiet, and K.M. Pryer. 2016. A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data. Taxon 65: 708–722. Reprint PDF

Pryer, K.M., L. Huiet, F.-W. Li, C.J. Rothfels, and E. Schuettpelz. 2016.
Maidenhair ferns, Adiantum, are indeed monophyletic and sister to shoestring ferns, vittarioids (Pteridaceae). Systematic Botany 41: 17-23. Reprint PDF

2011 – 2015

Kao, T.-T., K.M. Pryer, M.D. Turner, R.A. White, and P. Korall. 2015. Origins of the endemic scaly tree ferns on the Galápagos and Cocos Islands. International Journal of Plant Sciences 176: 869-879.  Reprint PDF

Grusz, A.L. and K.M. Pryer. 2015. Development of microsatellite markers for the apomictic triploid fern, Myriopteris lindheimeri (Pteridaceae). Applications in Plant Sciences 3. doi:10.3732/apps.1500061. Reprint PDF

Schuettpelz, E., K.M. Pryer, and M.D. Windham. 2015. A unified approach to taxonomic delimitation in the fern genus Pentagramma (Pteridaceae). Systematic Botany 40: 629-644. Reprint PDF

Wolf, P.G., E.B. Sessa, D.B. Marchant, F.-W. Li, C.J. Rothfels, E.M. Sigel, M.A. Gitzendanner, C.J. Visger, J.A. Banks, D.E. Soltis, P.S. Soltis, K.M. Pryer, and J.P. Der.  2015. An exploration into fern genome space. Genome Biology and Evolution 7: 2533-2544. Reprint PDF

Li, F.-W., C.J. Rothfels, M. Melkonian, J.C. Villarreal, D.W. Stevenson, S.W. Graham, G.K.-S. Wong, S. Mathews, and K.M. Pryer. 2015. The origin and evolution of phototropins. Frontiers in Plant Science 6: 637. Reli-et-al-2015-frontiersprint PDF

Li, F.-W., M. Melkonian, C.J. Rothfels, J.C. Villareal, D.W. Stevenson, S.W. Graham, G.K.-S. Wong, K.M. Pryer, and S. Mathews.  2015Phytochrome diversity in green plants and the origin of canonical plant phytochromes. Nature Communications 6: 7852. Reprint PDF

MEDIA COVERAGEFaculty 1000DukeTodayScience

Rothfels C.J., F.-W. Li, E.M Sigel., L. Huiet, A. Larsson, D.O. Burge, M. Ruhsam, M. Deyholos, D. Soltis, N. Stewart, S. Shaw, L.M. Pokorny, T. Chen, C. dePamphilis, L. DeGironimo, D.W. Stevenson, S.W. Graham, G.K.-S. Wong, and K.M. Pryer. 2015. The evolutionary history of ferns inferred from 25 low-copy nuclear genes. American Journal of Botany 102: 1089-1107. Reprint PDF

Huiet, L., M. Lenz, J.K. Nelson, K.M. Pryer, and A.R. Smith. 2015. Adiantum shastense, a new species of maidenhair fern from California. Phytokeys 53:  73-81. Reprint PDF

Rothfels C.J., A.K. Johnson, P.H. Hovenkamp, D.L. Swofford, H.C. Roskam, C.R. Fraser-Jenkins, M.D. Windham, and K.M. Pryer. 2015. Natural hybridization between genera that diverged from each other approximately 60 million years ago. American Naturalist 185: 433-442. Reprint PDF

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Rothfels C.J., A.K. Johnson, M.D. Windham, and K.M. Pryer. 2014. Low-copy nuclear data confirm rampant allopolyploidy in the Cystopteridaceae (Polypodiales). Taxon 63: 1026-1036. Reprint PDF

Li, F.-W. and K.M. Pryer. 2014. Crowdfunding the Azolla fern genome project: a grassroots approach. GigaScience 3: 16.  Reprint PDF

Sessa, E.B., J.A. Banks, M.S. Barker, J.P. Der, A.M. Duffy, S.W. Graham, M. Hasebe, J. Langdale, F.-W. Li, D.B. Marchant, K.M. Pryer, C.J. Rothfels, S.J. Roux, M.L. Salmi, E.M. Sigel, D.E. Soltis, P.S. Soltis, D.W. Stevenson, and P.G. Wolf. 2014. Between two fern genomes. GigaScience 3: 15.  Reprint PDF

Pryer, K.M. 2014. “This lowly fern has massive green potential”. The Toronto Globe and Mail 30 May 2014; “Offering an ancient solution to a modern problem”. Contra Costa Times 1 June 2014. [Published Op-Ed].

Sigel, E.M., M.D. Windham, and K.M. Pryer. 2014. Evidence for reciprocal origins in Polypodium hesperium (Polypodiaceae): A natural fern model system for investigating how multiple origins shape allopolyploid genomes. American Journal of Botany 101: 1476–1485. Reprint PDF

Li, F.-W., J.C. Villarreal, S. Kelly, C.J. Rothfels, M. Melkonian, E. Frangedakis, M. Ruhsam, E.M. Sigel, J.P. Der, J. Pittermann, D.O. Burge, L. Pokorny, A. Larsson, T. Chen, S. Weststrand, P. Thomas, E. Carpenter, Y. Zhang, Z. Tian, L. Chen, Z. Yan, Y. Zhu, X. Sun, J. Wang, D.W. Stevenson, B.J. Crandall-Stotler, A.J. Shaw, M.K. Deyholos, D.E. Soltis, S.W. Graham, M.D. Windham, J.A. Langdale, G. K.-S. Wong, S. Mathews, and K.M. Pryer. 2014. Horizontal transfer of an adaptive chimeric photoreceptor from bryophytes to ferns. Proceedings of the National Academy of Sciences, USA 111: 6672-6677. Reprint PDF

    MEDIA COVERAGENew York TimesThe EconomistCurrent BiologyNational GeographicCosmos MagazineDuke TodayScientific AmericanDecoded ScienceNature Reviews Genetics

Sigel, E.M., M.D. Windham, C.H. Haufler, and K.M. Pryer. 2014. Phylogeny, divergence time estimates, and phylogeography of the diploid species of the Polypodium vulgare complex. Systematic Botany 39: 1042-1055. Reprint PDF

Grusz, A.L., M.D. Windham, G. Yatskievych, L. Huiet, G.J. Gastony, and K.M. Pryer. 2014. Patterns of diversification in the xeric-adapted fern genus Myriopteris (Pteridaceae). Systematic Botany 39: 698-714. Reprint PDF.

Sigel, E.M., M.D. Windham, A.R. Smith, R.J. Dyer, and K.M. Pryer. 2014.  Rediscovery of Polypodium calirhiza (Polypodiaceae) in Mexico. Brittonia 66: 278-286. Reprint PDF

Korall, P. and K.M. Pryer. 2014. Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal. Journal of Biogeography 41: 402-413. Reprint PDF

Rothfels C.J., A. Larsson, F.-W. Li, E.M. Sigel, L. Huiet, D.O. Burge, M. Ruhsam, S.W. Graham, D. Stevenson, G.K.-S. Wong, P. Korall, and K.M. Pryer. 2013. Transcriptome-mining for single-copy nuclear markers in ferns. PLoS ONE 8(10): e76957. doi:10:1371/journal.pone.0076957.  Reprint PDF

León, B., C.J. Rothfels, M. Arakaki, K.R. Young, and K.M. Pryer. 2013. Revealing a cryptic fern distribution through DNA sequencing: Pityrogramma trifoliata in the western Andes of Peru. American Fern Journal 103. 40-48. Reprint PDF

Wang, Z.R. and K.M. Pryer. 2013. Gymnocarpium. Pp. 257-259 in 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. Reprint PDF

Wang, Z.R., C.H. Haufler, K.M. Pryer, and M. Kato. 2013. Cystopteridaceae. Pp. 257 in 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. Reprint PDF

Rothfels, C.J., M.D. Windham, and K.M. Pryer. 2013. A plastid phylogeny of the cosmopolitan fern family Cystopteridaceae (Polypodiopsida). Systematic Botany 38: 295–306. Reprint PDF

Pryer, K.M. 2012. The distinguished legacy of DMB: Donald MacPhail Britton (1923-2012). American Fern Journal 102: 241-251. Reprint PDF

Beck, J.B., J. Allison, K.M. Pryer, and M.D. Windham. 2012. Identifying multiple origins of polyploid taxa: a multilocus study of the hybrid cloak fern (Astrolepis integerrima; Pteridaceae). American Journal of Botany 99: 1857-1865. Reprint PDF

Li, F.-W., K.M. Pryer, and M.D. Windham. 2012. Gaga, a new fern genus segregated from Cheilanthes (Pteridaceae). Systematic Botany 37: 845-860. Reprint PDF

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Johnson, A.K., C.J. Rothfels, M.D. Windham, and K.M. Pryer. 2012. Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae). American Journal of Botany 99: 1118-1124. Reprint PDF

Rothfels, C.J.*, M.A. Sundue*, L.-Y. Kuo, A. Larsson, M. Kato, E. Schuettpelz, and K.M. Pryer. 2012. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61: 515-533. *equally contributing. Reprint PDF

Rothfels, C.J.*, A. Larsson*, L.-Y. Kuo*, P. Korall, W.-L. Chiou, and K.M. Pryer.2012. Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Systematic Biology 61: 490-509. doi: 10.1093/sysbio/sys001. *equally contributing. Reprint PDF (Cover)

Li, F.-W., L.-Y. Kuo, C.J. Rothfels, A. Ebihara, W.-L. Chiou, M.D. Windham, and K.M. Pryer. 2011. rbcL and matK earn two thumbs up as the core DNA barcode for ferns. PLoS One 6: e26597. Reprint PDF

Beck, J.B., M.D. Windham, and K.M. Pryer. 2011. Do asexual lineages lead short evolutionary lives? A case-study from the fern genus AstrolepisEvolution 65: 3217-3229. Reprint PDF

Sigel, E.M., M.D. Windham, L. Huiet, G. Yatskievych, and K.M. Pryer. 2011. Species relationships and farina evolution in the cheilanthoid fern genus Argyrochosma (Pteridaceae). Systematic Botany 36: 554-564. Reprint PDF

Wolf, P.G., J.P. Der, A.M. Duffy, J.B. Davidson, A.L. Grusz, and K.M. Pryer. 2011. The evolution of chloroplast genes and genomes in ferns. Plant Molecular Biology 76: 251–261. Reprint PDF

2000 – 2010

Pryer, K.M., E. Schuettpelz, L. Huiet, A.L. Grusz, C.J. Rothfels, T. Avent, D. Schwartz, and M.D. Windham. 2010. DNA barcoding exposes a case of mistaken identity in the fern horticultural trade. Molecular Ecology Resources 10: 979-985. Reprint PDF

Korall, P., E. Schuettpelz, and K.M. Pryer. 2010. Abrupt deceleration of molecular evolution linked to the origin of arborescence in ferns. Evolution 64: 2786-2792. Reprint PDF

Beck, J.B., M.D. Windham, G. Yatskievych, and K.M. Pryer. 2010. A diploids-first approach to species delimitation and interpreting polyploid evolution in the fern genus Astrolepis (Pteridaceae). Systematic Botany 35: 223-234. Reprint PDF

Schneider, H., A.R. Smith, and K.M. Pryer. 2009. Is morphology really at odds with molecules in estimating fern phylogeny? Systematic Botany 34: 455-475. Reprint PDF

Windham, M.D., L. Huiet, E. Schuettpelz, A.L. Grusz, C. Rothfels, J. Beck, G. Yatskievych, and K.M. Pryer. 2009. Using plastid and nuclear DNA sequences to redraw generic boundaries and demystify species complexes in cheilanthoid ferns. American Fern Journal 99: 128-132. Reprint PDF

Grusz, A.L., M.D. Windham, and K.M. Pryer. 2009. Deciphering the origins of apomictic polyploids in the Cheilanthes yavapensis complex (Pteridaceae). American Journal of Botany 96: 1636-1645. Reprint PDF

Schuettpelz, E. and K.M. Pryer. 2009. Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy. Proceedings of the National Academy of Sciences, USA 106: 11200-11205 (cover). Reprint PDF  Cover

Pryer, K.M. and D.J. Hearn. 2009. Evolution of leaf form in marsileaceous ferns: evidence for heterochrony. Evolution 63: 498-513. Reprint PDF

Pryer, K.M. and E. Schuettpelz. 2009. Ferns. Chapter 14 in S.B. Hedges and S. Kumar (eds.) Timetree of Life. Oxford University Press, New York. pp. 153-156.  Reprint PDF

Schuettpelz, E., A.L. Grusz, M.D. Windham, and K.M. Pryer. 2008. The utility of nuclear gapCp in resolving polyploid fern origins. Systematic Botany 33: 621-629. Reprint PDF

Hennequin, S., E. Schuettpelz, K.M. Pryer, A. Ebihara, and J.-Y. Dubuisson. 2008. Divergence times and the evolution of epiphytism in filmy ferns (Hymenophyllaceae) revisited. International Journal of Plant Sciences 169: 1278-1287. Reprint PDF

Nagalingum, N.S., M.D. Nowak, and K.M. Pryer. 2008. Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and SalviniaBotanical Journal of the Linnaean Society 157: 673-685. Reprint PDF

Rothfels, C.J., M.D. Windham, A.L. Grusz, G.J. Gastony, and K.M. Pryer. 2008. Toward a monophyletic Notholaena (Pteridaceae): Resolving patterns of evolutionary convergence in xeric-adapted ferns. Taxon 57: 712-724. Reprint PDF

Schuettpelz, E. and K.M. Pryer. 2008. Fern Phylogeny. Chapter 15 in Ranker, T.A. and C.H. Haufler (eds.). The Biology and Evolution of Ferns and Lycophytes. Cambridge Univ. Press., pp. 395-416. Reprint PDF

Smith A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. Wolf. 2008. Fern Classification. Chapter 16 in Ranker, T.A. and C.H. Haufler (eds.). The Biology and Evolution of Ferns and Lycophytes. Cambridge Univ. Press., pp. 417-467. Reprint PDF

Metzgar, J.S., J.E. Skog, E.A. Zimmer, and K.M. Pryer. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany 33: 31-36. Reprint PDF

Christenhusz, M.J.M., H. Tuomisto, J.S. Metzgar, and K.M. Pryer. 2008. Evolutionary relationships within the Neotropical, eusporangiate fern genus Danaea (Marattiaceae). Molecular Phylogenetics and Evolution 46: 34-48. Reprint PDF

Schuettpelz E. and K.M. Pryer. 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037-1050. Reprint PDF

Haugen, P., D. Bhattacharya, J.D. Palmer, S. Turner, L.A. Lewis, and K.M. Pryer.2007. Cyanobacterial ribosomal RNA genes with multiple, endonuclease-encoding group I introns. BMC Evolutionary Biology 7: 159. Reprint PDF

Metzgar, J.S., H. Schneider, and K.M. Pryer. 2007. Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae). International Journal of Plant Sciences 168: 1045-1053. Reprint PDF

Schuettpelz E., H. Schneider, L. Huiet, M.D. Windham, and K.M. Pryer. 2007. A molecular phylogeny of the fern family Pteridaceae: assessing overall relationships and the affinities of previously unsampled genera. Molecular Phylogenetics and Evolution 44: 1172-1185. Reprint PDF

Korall, P., D.S. Conant, J.S. Metzgar, H. Schneider, and K.M. Pryer. 2007. A molecular phylogeny of scaly tree ferns (Cyatheaceae). American Journal of Botany 94: 873-886 (cover). Reprint PDF

Nagalingum, N.S., H. Schneider, and K.M. Pryer. 2007. Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus MarsileaSystematic Botany 32: 16-25. Reprint PDF

Schuettpelz, E., P. Korall, and K.M. Pryer. 2006. Plastid atpA data provide improved support for deep relationships among ferns. Taxon 55: 897-906. Reprint PDF

Schuettpelz, E. and K.M. Pryer. 2006. Reconciling extreme branch length differences: decoupling time and rate through the evolutionary history of filmy ferns. Systematic Biology 55: 485-502. [2006 Publisher’s Award for Excellence in Systematic Research]. Reprint PDF

Korall, P., K.M. Pryer, J. Metzgar, H. Schneider, and D.S. Conant. 2006. Tree ferns: monophyletic groups and their relationships as revealed by four protein-coding plastid loci. Molecular Phylogenetics and Evolution 39: 830-845. Reprint PDF

Nagalingum, N.S., H. Schneider, and K.M. Pryer. 2006. Comparative morphology of reproductive structures in heterosporous water ferns and a reevaluation of the sporocarp. International Journal of Plant Sciences 167: 805-815. Reprint PDF

Korall, P., D.S. Conant, H. Schneider, K. Ueda, H. Nishida, and K.M. Pryer. 2006. On the phylogenetic position of Cystodium: it’s not a tree fern – it’s a polypod! American Fern Journal 96: 45-53. Reprint PDF

Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. Wolf. 2006. A classification for extant ferns. Taxon 55: 705-731. Reprint PDF.  #1 of just 10 papers Google calls a “Classic paper” in Botany. For translation into Ukrainian click here

Wikström, N. and K.M. Pryer. 2005. Incongruence between primary sequence data and the distribution of a mitochondrial atp1 group II intron among ferns and horsetails. Molecular Phylogenetics and Evolution 36: 484-493. Reprint PDF

Schneider, H., E. Schuettpelz, K.M. Pryer, R. Cranfill, S. Magallón, R. Lupia. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553-557. Reprint PDF

Pryer, K.M., H. Schneider and S. Magallón. 2004. The radiation of vascular plants. In J. Cracraft and M. J. Donoghue, eds. Assembling the Tree of Life, 138-153. Oxford University Press, London. Reprint PDF

Pryer, K.M., E. Schuettpelz, P.G. Wolf, H. Schneider, A.R. Smith, R. Cranfill. 2004. Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. American Journal of Botany 91: 1582-1598. [Special invited paper]  Reprint PDF

Des Marais, D.L., A.R. Smith, D.M. Britton, and K.M. Pryer. 2003. Phylogenetic relationships and evolution of extant horsetails,Equisetum, based on chloroplast DNA sequence data (rbcL andtrnL-F). International Journal of Plant Sciences 164: 737-751. Reprint PDF

Dubuisson, J.-Y., S. Hennequin, E.J.P. Douzery, R. Cranfill, A.R. Smith and K.M. Pryer. 2003. rbcL phylogeny of the fern genusTrichomanes (Hymenophyllaceae), with special reference to neotropical taxa. International Journal of Plant Sciences 164: 753-761. Reprint PDF

Pryer, K.M., H. Schneider, E.A. Zimmer and J.A. Banks. 2002. Deciding among green plants for whole genome studies. Trends in Plant Science 7: 550-554. Reprint PDF

Schneider, H. and K.M. Pryer. 2002. Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. International Journal of Plant Sciences 163: 485-505 (cover). Reprint PDF

Schneider, H., K.M. Pryer, R. Cranfill, A.R. Smith and P.G. Wolf. 2002. Evolution of vascular plant body plans: a phylogenetic perspective. Pp. 330-364 in Developmental Genetics and Plant Evolution, Q.C.B. Cronk, R.M. Bateman, and J.A. Harris, eds. Taylor and Francis, Philadelphia. Reprint PDF

Smith, A.R., H. Tuomisto, K.M. Pryer, J.S. Hunt and P.G. Wolf. 2001. Metaxya lanosa, a second species in the genus and fern family Metaxyaceae. Systematic Botany 26: 480-486. Reprint PDF

Pryer, K.M., A.R. Smith, J.S. Hunt, and J.-Y.Dubuisson. 2001. rbcLdata reveal two monophyletic groups of filmy ferns (Filicopsida:Hymenophyllaceae). American Journal of Botany 88: 1118-1130 (cover). Reprint PDF

Pryer, K.M., H. Schneider, A.R. Smith, R. Cranfill, P.G. Wolf, J.S. Hunt and S.D. Sipes. 2001. Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Nature 409: 618-622 (cover). Reprint PDF

Lupia, R., H. Schneider, G.M. Moeser, K.M. Pryer and P.R. Crane. 2000. Marsileaceae sporocarps and spores from the Late Cretaceous of Georgia, U.S.A.International Journal of Plant Sciences 161: 975-988. Reprint PDF

Prior to 2000

Wolf, P.G., S.D. Sipes, M.R. White, M.L. Martines, K.M. Pryer, A.R. Smith and K. Ueda. 1999. Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: evidence from rbcL nucleotide sequences. Plant Systematics and Evolution 219: 263-270. Reprint PDF

Turner, S., K.M. Pryer, V.P.W. Miao and J.D. Palmer. 1999. Investigating deep phylogenetic relationships among cyanobacteria and plastids by small subunit rRNA sequence analysis. Journal of Eukaryotic Microbiology 46: 327-338. Reprint PDF

Pryer, K.M. 1999. Phylogeny of marsileaceous ferns and relationships of the fossil Hydropteris pinnata reconsidered. International Journal of Plant Sciences 160: 931-954 (cover). Reprint PDF

Wolf, P.G., K.M. Pryer, A.R. Smith and M. Hasebe. 1998. Phylogenetic studies of extant pteridophytes. Pp. 541-556 in D.E. Soltis, P.S. Soltis and J.J. Doyle, eds. Molecular systematics of plants II: DNA sequencing. Kluwer Academic Publishers, Boston. Reprint PDF

Pryer, K.M., A.R. Smith and J.E. Skog. 1995. Phylogenetic relationships of extant ferns based on evidence from morphology and rbcL sequences. American Fern Journal 85 (Special Issue): 205-282. Reprint PDF

Hasebe, M., P.G. Wolf, K.M. Pryer, K. Ueda, M. Ito, R. Sano, G.J. Gastony, J. Yokoyama, J.R. Manhart, N. Murakami, E.H. Crane, C.H. Haufler and W.D. Hauk.1995. Fern phylogeny based on rbcLnucleotide sequences. American Fern Journal 85 (Special Issue): 134-181. Reprint PDF

Pryer, K.M. and C.H. Haufler. 1993. Isozymic and chromosomal evidence for the allotetraploid origin of Gymnocarpium dryopteris(Dryopteridaceae). Systematic Botany18: 150-172. Reprint PDF

Pryer, K.M. 1993. Gymnocarpium. Pp. 258-262 in Flora of North America North of Mexico, Vol.2. Flora of North America Editorial Committee (ed.). Oxford University Press, New York. Reprint PDF

Pryer, K.M. 1992. The status of Gymnocarpium heterosporum and G. robertianum in Pennsylvania. American Fern Journal 81: 34-39. Reprint PDF

Pryer, K.M. 1990. The limestone oak fern: new to Manitoba. The Blue Jay 48:192-195. Reprint PDF

Pryer, K.M. and L.R. Phillippe. 1989. A synopsis of the genus Sanicula in eastern Canada. Canadian Journal of Botany 67: 694-707. Reprint PDF

McNeill, J. and K.M. Pryer. 1985. The status and typification of Phegopteris and GymnocarpiumTaxon 34: 136-156. Reprint PDF

Pryer, K.M., D.M. Britton and J. McNeill. 1983. A numerical analysis of chromatographic profiles in North American taxa of the fern genus GymnocarpiumCanadian Journal of Botany 61: 2592-2602. Reprint PDF

Pryer, K.M. and D.M. Britton. 1983. Spore studies in the genus GymnocarpiumCanadian Journal of Botany 61: 377-388. Reprint PDF

Sarvela, J., D.M. Britton and K.M. Pryer. 1981. Studies on the Gymnocarpium robertianum complex in North America. Rhodora 83: 421-431. Reprint PDF

Chase, R., K.M. Pryer, R. Baker and D.M. Madison. 1978. Responses to conspecific chemical stimuli in the terrestrial snail Achatina fulica (Pulmonata: Sigmuretha). Behavioural Biology 22: 302-315. Reprint PDF


Argus, G.W. & K.M. Pryer. 1990. Rare Vascular Plants in Canada – Our Natural Heritage. Canadian Museum of Nature, Ottawa. 277 pp.

Pryer, K.M. & G.W. Argus (eds.). 1987. Atlas of the Rare Vascular Plants of Ontario. Part 4. National Museum of Natural Sciences, Ottawa. 472 pp.

Electronic Publications

Pryer, Kathleen M., Alan R. Smith, and Carl Rothfels. 2008. Polypodiidae Cronquist, Takht. & Zimmerm. 1966. Leptosporangiate Ferns. Version 23 December 2008. in The Tree of Life Web Project,

Pryer, Kathleen M., Alan R. Smith, and Carl Rothfels. 2009. Polypodiopsida Cronquist, Takht. & Zimmerm. 1966. Ferns. Version 14 January 2009. in The Tree of Life Web Project,