Kinosian, S.P and P.G. Wolf. 2022. The natural history of model organisms: The biology of the C. richardii as a tool for understanding plant evolution. eLife 2022;11:e75019

Kinosian, S.P., C.A. Rowe, P.G. Wolf. 2022. Why do heterosporous plants have so few chromosomes? Frontiers in Plant Science 13:807302. doi: 10.3389/fpls.2022.807302.

Koontz A, W.D. Pearse, P.G. Wolf. 2022. Pronounced genetic separation among varieties of the Primula cusickiana species complex, a Great Basin endemic. Systematic Botany In Press


Wolf, P. G. 2021. Cellular Processes: Chromosome number pattern seen in ferns. Open Access Government Jan 2021: 234-235.


Rowe CA, Lichvar RW and Wolf PG (2020) How Many Tree Species of Birch Are in Alaska? Implications for Wetland Designations. Frontiers in Plant Science 11:750. doi: 10.3389/fpls.2020.00750

Kinosian SP, WD Pearse, and PG Wolf. 2020.There and back again: Reticulate evolution in Ceratopteris. American Fern Journal 110: 193 – 210.

Kinosian SP, WD Pearse, and PG Wolf. 2020. Cryptic diversity in the model fern genus Ceratopteris (Pteridaceae). Molecular Phylogenetics and Evolution 152: 106938.


Marchant DB, EB Sessa, PG Wolf, K Heo, WB Barbazuk, PS Soltis, DE Soltis. 2019. The C-Fern (Ceratopteris richardii) genome: insights into plant genome evolution with the first partial homosporous fern genome assembly. Scientific Reports 9: 18181.

Wolf PG, Rowe CA, Kinosian SP, Der JP, Lockhart PJ, Shepherd LD, McLenachan PA, Thomson JA. 2019. Worldwide relationships in the fern genus Pteridium (bracken) based on nuclear genome markers. American Journal of Botany 106(10): 1365–1376

Rowe CA, PG Wolf, RW Lichvar. 2019. Do genetic differences explain the ability of an alkaline shrub to grow in both uplands and wetlands? Western North American Naturalist: 79: 260–269

Wolf, PG and MS Barker. 2019. Current status and future prospects for fern and lycophyte genomics: introduction to an American Fern Journal special issue. American Fern Journal 109 (3): 177-182

Robison TA and PG Wolf. 2019. ReFernment: an R package for annotating RNA editing in plastid genomes. Applications in Plant Sciences 7(2): e1216


Robison TA, Grusz AL, Wolf PG, Mower JP, Fauskee BD, Sosa K, Schuettpelz E. 2018. Mobile elements shape plastome evolution in ferns. Genome Biology and Evolution 10: 2558–2571

Lemon, JB and PG Wolf. 2018. Genetic differentiation between endemic Eriogonum soredium and its common relative E. shockleyi (Polygonaceae). Systematic Botany 43: 901-909

Ellis, MW, PG Wolf, CA Rowe. 2018. Field identification of Eriogonum corymbosum vars. nilesii and aureum (Polygonaceae). Phytotaxa 382: 293–296

Rowe, CA, Hauber, DP, Wolf PG. 2018. Genomic variation of introduced Salvinia minima in southeastern United States. Aquatic Botany 151: 38-42

Li, F-W et al, TA Robison, PG Wolf. 2018. Fern genomes elucidate land plant evolution and cyanobacterial symbioses. Nature Plants 4: 460–472.

Schilling MP, Gompert Z, Li F-W, Windham MD, Wolf PG. 2018. Admixture, evolution, and variation in reproductive isolation in the Boechera puberula clade. BMC Evolutionary Biology 18: 61

Wolf, PG, Robison TA, Johnson MG, Sundue MA, Testo WL, Rothfels CJ. 2018. Target Sequence Capture of Nuclear-Encoded Genes for Phylogenetic Analysis in Ferns. Applications in Plant Sciences 6(5): e1148


Cai S, G Chen, Y Wang, Y Huang, DB Marchant, Y Wang, Q Yang, F Dai, A Hills, PJ Franks, E Nevo, DE Soltis, PS Soltis, E Sessa, PG Wolf, D Xue, G Zhang, BJ Pogson, MR Blatt, Z-H Chen. 2017. Evolutionary conservation of ABA signaling for stomatal closure in ferns. Plant Physiology 174: 732-747. doi:10.1104/pp.16.01848


The Pteridophyte Phylogeny Group I. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution. 54: 563–603.

Longhurst ML, DC Coster, PG Wolf, AM Duffy, H Lee, DT Campbell. 2016. Multi-year professional development grounded in educative curriculum focused on integrating technology with reformed science teaching principles. School Science and Mathematics 116: 430-441


Wolf, PG and Lichvar, RW. Video series on fern biology: 2a) Natural History of Ferns 2b) Identifying Ferns. URL:

Wolf PG, Rowe CA, Der JP, Schilling MP, Visger CJ, Thomson J. 2015. Origins and diversity of a cosmopolitan fern genus on an island archipelago. AoB Plants 7: plv118; DOI: 10.1093/aobpla/plv118

Wolf PG, Sessa EB, Marchant DB, Li F-W, Rothfels CJ, Sigel EM, Gitzendanner MA, Visger CJ, Banks JA, Soltis DE, Soltis PS, Pryer KM, Der JP. 2015. An exploration into fern genome space. Genome Biology and Evolution 7: 2533-2544. DOI: 10.1093/gbe/evv163

Ellis M.W., P.G. Wolf, S. Bardot, J.A. Walton, C.A. Rowe, S. Kulpa, and KA. Mock. 2015. Further Examination of the Geographic Range of Eriogonum corymbosum var. nilesii (Polygonaceae). Phytotaxa 203: 279–286


Schilling, M.P., Wolf, P. G., Duffy, A. M., Rai, H. S., Richardson, B. A., Mock, K. E. 2014. Genotyping-by-sequencing for Populus population genomics: an assessment of genome sampling patterns and filtering approaches. PLoS One 9(4): e95292.

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.

Duffy AM, MC Stensvold, DR Farrar. 2014. Independent gametophytes of Hymenophyllum wrightii in North America: not as rare as we thought. American Fern Journal 105: 45–55.

Campbell, T. D., B. A. ZuWallack, D.T, M. Longhurst, B. Shelton, P. G. Wolf. 2014. An examination of the changes in science teaching orientations and technology-enhanced tools for student learning in the context of professional development. International Journal of Science Education 36: 1815-1848.

Davidson, J. B. S. L. Durham, and P. G. Wolf. 2014. Breeding system of the threatened endemic, Primula cusickiana var. maguirei (Primulaceae). Plant Species Biology 29: e55–e63.


Rai, H. S., K. E. Mock, B. A. Richardson, R. C. Cronn, K. J. Jayden, J. W. Wright, B. J. Knaus, P. G. Wolf. 2013. Transcriptome characterization and detection of gene expression differences in aspen (Populus tremuloides). Tree Genetics and Genomes 9: 1031-1041. DOI: 10.1007/s11295-013-0615-y.

Mock, K. E., B. A. Richardson, and P. G. Wolf. 2013. Molecular tools and aspen management: a primer and prospectus. Forest Ecology and Management 299: 6-13

Duffy, A. M., Wolf, P. G., Barrow, J., Longhurst, M., Campbell, D.T. & Beckett, H. 2013. Ecological investigations within an interactive plant community simulation. Science Scope 36(8): 42-51.

Campbell, D.T., Longhurst, M., Duffy, A. M, Wolf, P. G., & Shelton, B. E. 2013. Science teaching orientations and technology-enhanced tools for student learning in science. Research in Science Education. 43:2035–2057


Campbell, D.T., Longhurst, M., Duffy, A. M., Wolf, P. G., & Nagy, R. 2012. Investigating human impact in the environment with faded scaffolded inquiry supported by technologies. Science Activities. 49: 4

Mock, K.E., Callahan, C.M., Islam-Faridi, M.N., Shaw, J.D., Rai, H.S., Sanderson, S.C., Rowe, C.A., Ryel, R.J., Madritch, M.D., Gardner, R.S., Wolf, P.G., 2012. Widespread triploidy in western North American aspen (Populus tremuloides). PLoS ONE 7(10): e48406. doi:10.1371/journal.pone.0048406

Wolf, P. G. and K. G. Karol. 2012. Plastomes of bryophytes, lycophytes and ferns. Pages 89 – 102 In Advances in Photosynthesis and Respiration. Vol 35. “Genomics of chloroplasts and mitochondria”. Editors: Ralph Bock & Volker Knoop. Springer, Dordrecht.

Wolf, P. G. 2012. Plastid genome diversity. Pages 145-154. In “Plant Genome Diversity” by Edited by Johann Greilhuber, Jonathan Wendel, Ilia J. Leitch, Jaroslav Dolezel. Springer Verlag.

Campbell, D. T, P. G. Wolf, J. P. Der, E. Packenham, and N. Abd-Hamid. 2012. Scientific inquiry in the genetics laboratory: biologists and university science teacher educators collaborating to increase engagement in science process. Journal of College Science Teaching 42: 82-89


Davidson, J. B. and P. G. Wolf. 2011. Natural history of Maguire primrose, Primula cusickiana var. maguirei (Primulaceae). Western North American Naturalist 71: 327 – 337.

Banks, J. A., T. Nishiyama, M. Hasebe, J. L. Bowman, M. Gribskov, C. dePamphilis, P. G. Wolf et al. 2011. The Selaginella genome identifies changes in gene content associated with the evolution of vascular plants. Science 332: 960-963. DOI: 10.1126/science.1203810.

Der, J. P., M. S. Barker , N. J. Wickett, C. W. dePamphilis, P. G. Wolf. 2011. De novo characterization of the gametophyte transcriptome in bracken fern, Pteridium aquilinum. BMC Genomics 12: 99. doi:10.1186/1471-2164-12-99.

Wolf P.G., Der J.P., Duffy A.M., Davidson J.B., Grusz A.L., Pryer K.M. 2011. The evolution of chloroplast genes and genomes in ferns. Plant Molecular Biology 76: 251-261. DOI 10.1007/s11103-010-9706-4.


Karol, K. G., Arumuganathan, K., Boore, J. L., Duffy, A. M., Everett, K. D. E., Hall, J. D., &Hansen, S. K., Kuehl, J. V., Mandoli, D. F., Mishler, B. D., Olmstead, R. G., Renzaglia, K. S., Wolf, P. G. 2010. Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages. BMC Evol. Biol. 10: 321.

Campbell, D. T., S. W. Wang, H‐Y. Hsu, A. M. Duffy, and P. G. Wolf. 2010. Learning with Web Tools, Simulations, and Other Technologies in Science Classrooms. Journal of Science Education and Technology 19:505-511.

Wolf, P. G., Roper, J. M., Duffy, A. M. 2010. The evolution of chloroplast genome structure in ferns. Genome 53: 731-738.

Oliver, M. J., A. G. Murdock, B. D. Mishler, J. V. Kuehl, J. L. Boore, D. F. Mandoli, K. D. E. Everett, P. G. Wolf, *A. M. Duffy, K. G. Karol. 2010. Chloroplast genome sequence of the moss Tortula ruralis: gene content, polymorphism, and structural arrangement relative to other green plant chloroplast genomes. BMC Genomics 2010, 11:143

Barker, M. S. and P. G. Wolf. 2010. Unfurling fern biology in the genomics age. BioScience 60(3): 177-185.

Ellis, M. W. and P. G. Wolf. 2010. Teaching Species. Evolution: Education and Outreach 3: 89-98.


Wolf, P. G., A. M. Duffy, and J.M. Roper. 2009. Phylogenetic use of inversions in fern chloroplast genomes. American Fern Journal 99:132-133.

Ellis, M. W., J. M. Roper, &R. Gainer, J. P. Der, and P. G. Wolf. 2009. The taxonomic designation of Eriogonum corymbosum var. nilesii (Polygonaceae) is supported by AFLP and cpDNA analyses. Systematic Botany 34: 693-703.

Duffy, A. M., S. A. Kelchner, and P. G. Wolf. 2009. Conservation of selection on matK following an ancient loss of its flanking intron. Gene 438: 17-25.

Der, J. P., J. A. Thomson, J. K. Stratford & P. G. Wolf. 2009. Global chloroplast phylogeny and biogeography of bracken (Pteridium: Dennstaedtiaceae). American Journal of Botany 96: 1041-1049.


Bjerregaard, L. and P. G. Wolf. 2008. Strong genetic differentiation among neighboring populations of a locally endemic primrose. Western North American Naturalist 68: 66-75.

Wickett, N. J. , Y. Zhang, &S. K. Hansen, J. M. Roper, J. V. Kuehl, S. A. Plock, B. Goffinet, P. G. Wolf, C. W. dePamphilis, Jeffrey L. Boore. 2008. Functional gene losses occur with minimal size reduction in the plastid genome of the parasitic liverwort Aneura mirabilis. Molecular Biology and Evolution 25: 393–401.

Wolf, P.G. and J. M. Roper. 2008. Structure and evolution of fern plastid genomes. In: “Biology and Evolution of Ferns and Lycophytes” Edited by Tom A. Ranker and Christopher H. Haufler (Cambridge University Press)

Smith, A. R., K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, & P. G. Wolf. 2008. Fern Classification. In: “Biology and Evolution of Ferns and Lycophytes” Edited by Tom A. Ranker and Christopher H. Haufler (Cambridge University Press)


Roper, J.M, &S.K. Hansen, P.G. Wolf , K.G. Karol KG, D.F. Mandoli, K.D.E. Everett, J. Kuehl, J.L. Boore. 2007. The Complete Plastid Genome Sequence of Angiopteris evecta (G. Forst.) Hoffm. (Marattiaceae). American Fern Journal 97 (2): 95-106


Smith, A. R., K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider & P. G. Wolf. 2006. A classification for extant ferns. Taxon 55 (3): 705-731


Wolf, P.G, K.G. Karol, D. F. Mandoli, J. Kuehl, K. Arumuganathan, M. W. Ellis, B. D. Mishler, D. G. Kelch, R. G. Olmstead, J. L. Boore. 2005. The first complete chloroplast genome sequence of a lycophyte, Huperzia lucidula (Lycopodiaceae). Gene 350: 117-128.


Wolf, P. G., C. A. Rowe, M. Hasebe. 2004. High levels of RNA editing in a vascular plant chloroplast genome: analysis of transcripts from the fern Adiantum capillus-veneris. Gene 339: 89-97

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: 1557-1581

Nishiyama, T., M. Kugita, R. B. Sinclair, M. Sugita, C. Sugiura, T. Wakasugi, P. G. Wolf, K. Yamada, K. Yoshinaga, and M. Hasebe. 2004. Bryophytes are monophyletic and land plants comprise two extant lineages. Molecular Biology and Evolution, 21 (10): 1813-1819

Wolf, P. G., B. Doche, L. Gielly and P. Taberlet. 2004. Genetic structure of Rhododendron ferrugineum at a wide range of spatial scales. Journal of Heredity 95: 301-308


Wolf, P. G., C. A. Rowe, R. B. Sinclair, M. Hasebe. 2003. Complete nucleotide sequence of the chloroplast genome from a leptosporangiate fern, Adiantum capillus-veneris L. DNA Research 10 (2): 59-64.


Schneider, H., K. M. Pryer, R. Cranfill, A. R. Smith, P. G. Wolf. 2002. Phylogeny of vascular plants and the evolution of their body plans. In Developmental Genetics and Plant Evolution. Q. Cronk, R. M. Bateman, J. A. Hawkins (Eds.) Taylor and Francis, London.


Rowe, D. L., M. T. Murphy, R. C. Fleischer, and P. G. Wolf. 2001. High frequency of extra-pair paternity in Eastern Kingbirds. Condor 103: 845-851.

Sipes, S. D. and P. G. Wolf. 2001. Phylogenetic relationships within Diadasia, a group of specialist bees. Molecular Phylogenetics and Evolution 19: 144-156.

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

Archibald J.K., P.G. Wolf, V. J. Tepedino, and J. Bair. 2001. Genetic relationships and population structure of the endangered steamboat buckwheat, Eriogonum ovalifolium var. williamsiae (Polygonaceae). American Journal of Botany 88: 608-615.

Wolf, P. G., D. R. Campbell, N. M. Waser, *S. D. Sipes, T. R. Toler, and &J. K. Archibald. 2001. Tests of pre-and post-pollination barriers to hybridization between sympatric species of Ipomopsis (Polemoniaceae). American Journal of Botany 88: 213-219.

Wolf, P. G., H. Schneider, and T. A. Ranker. 2001. Geographic distributions of homosporous fern taxa: does dispersal obscure evidence of vicariance? Journal of Biogeography 28: 263-270.

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 relatives to seed plants. Nature 409: 618-622.


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

Soltis, P. S., D. E. Soltis, P. G. Wolf, D. L. Nickrent, S-M. Chaw, and R. L. Chapman. 1999. The Phylogeny of land plants inferred from 18S rDNA sequences: pushing the limits of rDNA signal? Molecular Biology and Evolution 16: 1774 – 1784.

1986 – 1998

See Paul’s CV under “Peer-Reviewed Journal Articles”