Peer-Reviewed publications from Villarreal lab (2015-on)

Members of the lab in bold

undergraduate* and master/PhDstudents/Post-docs** at the time of doing the research

For the entire publication list see


In review or in consideration

  1. Sierra, A.**, M. Alonso García**, C. Zartman & C. Villarreal A. The consequences of mating system and dispersal potential on the genetic structure of leaf-inhabiting bryophyte metapopulations in a fragmented Amazonian landscape. Submitted to Molecular Ecology. Preprint:
  2. Hernández-Rodríguez, E., C. Cerrejón, Y. Xiangbo, M. Noualhaguet, M. Indorf, C. Villarreal, N. Fenton.Undefined rareness patterns in inconspicuous boreal species. Submitted to the Forest Ecology and Management
  3. J.C. Villarreal, Meléndez O.*, Bethancourt R, Bethancourt A, Rodriguez-Castro L.**, López CA., Sedio BE, Saltonstall K. Two draft fungal genomes of leaf endophytes from tropical gymnosperms. Microbiology Resources Announcements.
  4. Gutierrez-Ortega, José Said & J.C. Villarreal. A possible case of adaptive radiation in cycads: A commentary on Zamia phylogeny and biogeography. Annals of Botany


Bell-Doyon, P.**,M.J. Mazerolle, L. Bélanger, Nicole Fenton, J.C. Villarreal A. 2024.  Differential impact of clearcut and insect outbreak on boreal lichens and bryophytes 50 years after disturbance. Biological Conservation. Bell-Doyon.etal.2024.Biolconservation

Sierra A.M.**, Meléndez O.*, Bethancourt R, Bethancourt A, Rodriguez-Castro L.**, López CA., Sedio BE, Saltonstall K, Villarreal JC.  2024. Leaf endophytes correlate with host metabolome expression in tropical gymnosperms. Journal of Chemical Ecology Sierra.etal.2024.CycadMetabolome

Hernández-Rodríguez, E., J.C. Villarreal, N. Fenton. In press. Patch level boreal bryophyte diversity driven by landscape heterogeneity Forest Ecology and Management Hernandez-Rodriguez_etal_ForestEcologyManagement_2024

Villarreal, J.C., N.B. Villarreal, L.F. De León. Panama says no to more mining – a win for environmentalists. Nature 625: 30. Villarreal-2024.Nature

Sierra P., A.**, S. Toupin, M. Alonso-García**, J.C. Villarreal A.2024. Diversity of symbiotic cyanobacteria in cycad coralloid roots using a short-read rbcL-X amplicon. Symbiosis. Sierra_et_al_Symbiosis_rbcLX


Bechteler, J., G. Peñaloza-Bojacá….27 authors, J.C. Villarreal A. 2023. Comprehensive phylogenomic time tree of bryophytes reveals deep relationships and uncovers gene incongruences in the last 500 million years of diversificationAmerican Journal of Botany 110 (11): e16249 Bechteler_etal_2023_AJB

Escolástico-Ortiz, D.A.**Charlotte Blasi, Jean-Philippe Bellenger, Nicolas Derome, & Juan Carlos Villarreal A. 2023. Differentially abundant bacteria drive the N2-fixation of a widespread moss in the forest-tundra transition zone. Symbiosis.pdf

Bell-Doyon, P.** 2023. Checklist of Lichens and Associated Fungi from Mingan Archipelago National Park Reserve, Québec, CanadaNortheastern Naturalist 30(3):304-328. Note: Further distribution or reproduction of the manuscript, either in whole or in part, except for personal research purposes, is prohibited without the written permission of the publisher, Eagle Hill Institutepdf.

Peñaloza-Bojacá, Gabriel F., Tiago Vilas-Boas, Juan C. Villarreal A, and Adaíses S. Maciel-Silva. 2023. “Differential Effects of Desiccation on Hornworts with Contrasting Life Histories in Tropical Montane Forests: A Functional Trait—Based Perspective” Forests 14, no. 2: 255.

Dennis Alejandro Escolástico-Ortiz,**, Lars Hedenäs, Dietmar Quandt, Dörte Harpke, Juan Larraín, Michael Stech & Juan Carlos Villarreal A. 2023. Cryptic speciation shapes the biogeographic history of a northern distributed moss. Botanical Journal of the Linen Society 201(1): 114-134, Escolastico-Ortiz.etal.2021. BJLinneanSociety.


Budke, J.M, M. Renner, J.C. Villarreal A., M. Stech (editors). 2022. A tribute to the contributions of Jeffrey Graham Duckett. Bryophyte Diversity and Evolution 45(1).

Cargill, D.C., S. Chantanaorrapint, R.-L. Zhu, R.-L., A. Asthana,L Li, K.S. Renzaglia, K.S. and J.C. Villarreal A. 2022. Resolving Relationships Within The Hornwort Genus Anthoceros. Bryophyte Diversity and Evolution 45(1): 26-43. Cargill.etal.2022.BDE

Peñaloza-Bojacá, G., A. Sierra**, H. Becher, K.S. Renzaglia, J.C. Villarreal A. 2022. HISTORICAL BIOGEOGRAPHY OF THE AUSTRAL HORNWORT GENUS PHAEOMEGACEROS (DENDROCEROTACEAE, ANTHOCEROTOPHYTA)Bryophyte Diversity and Evolution 45(1): 44-66. Peñaloza-Bojaca.etal.2022.BDE

Allen, N.S., G. Dauphin, G. J.C. Villarreal A. et al. 2022. Bryophytes Of Mangroves Of Bocas Del Toro, Panama  Bryophyte Diversity and Evolution 45(1): 133–150. Salazar_Allen.etal.2022.BDE

Bell-Doyon, P.**, V. Bellavance, L. Bélanger, M.J. Mazerolle, J.C. Villarreal A. 2022. Bacterial, Fungal, and Mycorrhizal Communities in the Soil Differ between Clearcuts and Insect Outbreaks in the Boreal Forest 50 Years after Disturbance. Forest Ecology and Management 523(1): 120493

Claudio M. Monteza-Moreno, Lilisbeth Rodriguez-Castro**, Pedro L. Castillo-Caballero, Edgar Toribio, Kristin Saltonstall. 2022. Arboreal camera trapping sheds light on seed dispersal of the world’s only epiphytic gymnosperm: Zamia pseudoparasitica. Ecology and Evolution 12(3): e8769. This paper has been highly covered by the press.

Marta Alonso-García**, Raquel Pino-Bodas, Juan Carlos Villarreal A. 2022. Co-dispersal of symbionts in the lichen Cladonia stellaris inferred from genomic data. Fungal Ecology. Alonso-Garcia.etal.FungalEcology.2022

Alonso García, M.** & J.C. Villarreal. 2022. Bacterial community of reindeer lichens differs between northern and southern lichen woodlands. Canadian Journal of Forest Research • 18 January 2022 • Alonso&Villarreal.2022 

Jairo Patiño, Irene Bisang, Bernard Goffinet, Lars Hedenäs, Stuart McDaniel, Silvia Pressel, Michael Stech, Claudine Ah-Peng, Ariel Bergamini, Richard T Caners, D Christine Cargill, Nils Cronberg, Jeffrey Duckett, Sarah Eppley, Nicole J Fenton, Kirsten Fisher, Juana González-Mancebo, Mitsuyasu Hasebe, Jochen Heinrichs, Kristoffer Hylander, Michael S Ignatov, Javier Martínez-Abaigar, Nagore G Medina, Rafael Medina, Dietmar Quandt, Stefan A Rensing, Karen Renzaglia, Matthew Renner, Rosa M Ros, Alfons Schäfer-Verwimp, Juan Carlos Villarreal, Alain Vanderpoorten. 2022. Unveiling the nature of a miniature world: a horizon scan of fundamental questions in bryology. Journal of Bryology.Patino_etal_2022_JBryology


Villarreal, J.C., B. Goffinet, N. Fenton, M. Favreau, S. Schuette & M. von Konrat. 2021.  Proccedings of Bryophytes, lichens, and northern ecosystems in a changing world (July 6-9, 2021). Bryological Times.

Rahmatpour, N., D.A. Hauser, J.M. Nelson, P.Y. Chen, J.C. Villarreal A., M.Y. Ho, F-W. Li. 2021. A novel thylakoid-less isolate fills a billion year gap in the evolution of Cyanobacteria. Current Biology

Villarreal, A., J.C.; M. Renaudin, A. Beaulieu-Laliberté* & J.P. Bellenger. 2021. Stigonema associated with boreal Stereocaulon possesses the alternative vanadium nitrogenase. The Lichenologist 53: 215–220. pdf

Bell-Doyon**, S.B. Selva & T. R. Mcmullin. 2021. Calicioid fungi and lichens from an unprotected intact forest ecosystem in Québec. Écosciences

Alonso, M.**, F. Grewe, S. Payette & J.C. Villarreal. 2021. Population genomics and genetic diversity of a reindeer lichen species in Eastern Canada Lichen Woodlands. American Journal of Botany

Jesse W. BreinholtSarah B. Carey..20 authorsJuan Carlos Villarreal, Evelyn Webb Williams & J. Gordon Burleigh.  2021. A target enrichment probe set for resolving the flagellate plant tree of life.  Applications in Plant Sciences


Frangedakis, E., M. Shimamura, J.C. Villarreal, F-W. Li, M. Tomaselli, M. Waller, K. Sakakibara, K. Renzaglia, P.Szövény 2020. The Hornworts: Morphology, evolution and development. The New Phytologist (Tansley Review)

Bouchard, R.*, G. Peñaloza-Boyacá**,  S. Toupin**,  Y. Guadalupe*, J. Gudiño, N. Salazar, F.-W.  Li & J. C. Villarreal A.  2020. Contrasting bacteriome of the hornwort Leiosporoceros dussii in two nearby sites with emphasis on the hornwort-cyanobacterial symbiosis. Symbiosis. pdf/
Bell-Doyon, P.**, &, Juan Carlos Villarreal A. 2020. New Notes on the Ecology of the Epiphytic gymnosperm and Panamanian endemic Zamia pseudoparasitica. Neotropical Naturalist 2: 1-7.Bell-Doyon&Villarreal.2020.NeotropicalNaturalist

Lavoie, C.*, M. Renaudin**, R. Troy McMullin, J. Gagnon, C. Roy, M.-E. Beaulieu, J. P. Bellenger & J. C. Villarreal A. 2020. Extremely low genetic diversity of Stigonema associated with Stereocaulon in eastern Canada. The Bryologist, 123(2):188-203: pdf

Alonso, M.**, J.C. Villarreal, K. McFarland & B. Goffinet. 2020. Population genomics and phylogeography of a clonal bryophyte with spatially separated sexes and extreme sex ratios. Frontiers in Plant Science.

T. Dawes*, J.C. Villarreal f, P. Szövény, I. Bisang, F-W. Li, D.A. Hauser, D. Quandt, D.C. Cargill & L.L. Forrest. 2020. Extremely low genetic diversity in the European clade of the model bryophyte Anthoceros agrestis Plant Systematics and Evolution. full paper here: Dawes_et_al-2020-Plant_Systematics_and_Evolution  

Li, F-W., T. Nishiyama, 24 authors…. J.C. Villarreal A., K. Sakakibara, P. Szövény. 2020. Anthoceros genomes illuminate the origin of land plants and the unique biology of hornworts. Nature plants 6, pages259–272.

Bell-Doyon, P.**, J. Laroche, K. Saltonstall, J.C. Villarreal A. 2020. Specialized bacteriome uncovered in the coralloid roots of the epiphytic gymnosperm, Zamia pseudoparasitica. Environmental DNA.

Mishler, B.D, D. C. Cargill, J. C. Villarreal, K. Renzaglia, B. Crandall-Stotler, and R. Stotler. 2020. Anthocerotae. Pp. 223–225 in Phylonyms: A Companion to the PhyloCode (K. de Queiroz, P. D. Cantino, and J. A. Gauthier, eds.). CRC Press, Boca Raton, FL.


OneKP Consortium. 2019One thousand plant transcriptomes and the phylogenomics of green plants. Nature.

Peñaloza, G.**, J.C. Villarreal A & A. Silva. 2019Phylogenetic and morphological circumscription of the genus Dendroceros Ness (Dendrocerotaceae; Anthocerotophyta), with the addition of two new subgenera. Systematics and Biodiversity. Penaloza_Bojaca_JCVA_Adaises_2019_Systematics&Biodiversity

David Bell, Wesley K. Gerelle, Qianshi Lin, Steve Joya, Ying Chang, Z. Nathan Taylor, Carl J Rothfels, Anders Larsson, Juan Carlos Villarreal, Fay-Wei Li, Lisa Pokorny, Peter Szövényi, Barbara Crandall-Stotler, Lisa deGeronimo, David Beerling, Michael Deyholos, Matt von Konrat, Shona Ellis, Tao Chen, Dennis Stevenson, Jeffrey D. Palmer and Sean W. Graham.  2019. Organellomic datasets confirm a cryptic consensus on (unrooted) land-plant relationships and provide new insights into bryophyte molecular evolution . American Journal of Botany. Bell_OrganellargenomicsBryophytes_2019_AJB

Nelson, J.M., D.A. Hauser, J.A. Gudiño, Y.A. Guadalupe*, J.C. Meeks, N. Salazar, J.C. Villarreal, F.-W. Li. 2019. Complete genomes of symbiotic cyanobacteria clarify the evolution of Vanadium-nitrogenase. Genome Biology and Evolution  11 ( 7): 1959–1964,

Anette Garrido, Jose Gudiño Ledezma, Armando A. Durant-Archibold, Noris Salazar Allen, Juan Carlos Villarreal A, and Mahabir P. Gupta. 2019. Chemical profiling of the gamethophyte and sporophyte from the Panamanian hornwort Leiosporoceros dussii (Leiosporocerotaceae) by HS-SPME-GC-MS. Natural Products Communications August: 1-4. Garrido.etal_.2019.NaturalProducts.pdf.

Têtu, B.,  & J.C Villarreal A., 2019. Bryophytes des murailles du Vieux-Québec. Carnets de bryologie 24 : 1-5.


Renzaglia, K.S., J.C. Villarreal A. & D. Garbary. 2018. Morphology supports the setaphyte hypothesis: mosses plus liverworts form a natural group. Bryophyte Diversity and Evolution 40(2): 11–17.

Sierra, A.M.**, J. Bechteler, D. Cardoso D, C. Zartman & J.C. Villarreal A. 2018. Divergence time analyses suggest a Miocene origin of the narrow Amazonian endemic rheophytic Ceratolejeunea temnantha (Spruce) Reiner-Drehwald (Porellales, Lejeuneaceae). Bryophyte Diversity and Evolution. 40(2): 55–67. Sierra_et_al_2018_bde00040p067

Villarreal, A., J.C., Monique Turmel; Maurane Bourgouin-Couture*, Jérôme Laroche, Noris Salazar-Allen, Fay-Wei Li, Shifeng Cheng, Karen Renzaglia & Claude Lemieux. 2018 Genome-wide organellar analyses from the hornwort Leiosporoceros dussii show low frequency of RNA editing. Plos One. Equal contribution, * undergraduate doi: /10.1371/journal.pone.0200491


Lewis, L.R., Ickert-Bond, S., Biersma, E.M, Convey, P., Goffinet, B., Hassel, K., Kruijer, K., La Farge, C., Metzgar, J., Stech, M., J.C. Villarreal , McDaniel, S.F. 2017. Future directions and priorities for Arctic bryophyte research. Artic Science 3: 475-497.

Lang, D., —J.C. Villarreal , –50 authors.2017. The P. patens chromosome-scale assembly reveals moss genome structure and evolutionlThe Plant Journal 93: 515–533.

Li, F.-W. f , J.C. Villarreal f,  Szövényi, P. 2017. Hornworts: an overlooked window into carbon-concentrating mechanism. Trends in Plant Science 22(4): 275-277. f  Equal contribution.

Villarreal, J.C ., Ducket, J.G., Pressel, S. 2017. Morphology, ultrastructure and phylogenetic affinities of the single-island endemic Anthoceros cristatus Steph.(Ascension Island). Journal of Bryology 39: .

Renzaglia, K.S., J.C. Villarreal,  Piatkowsky, B.T., Reagan, J.L. & Merced, A. 2017. Hornwort stomata: architecture and fate shared with 400 million year old fossil plant without leaves. Plant Physiology.


Long, D.G., Forrest, L.L., J.C. Villarreal & Crandall-Stotler. 2016.Taxonomic changes in Marchantiaceae, Corsiniaceae and Cleveaceae (Marchantiidae, Marchantiophyta). Phytotaxa 252(1): 077-080.

Villarreal, J.C., Crandall-Stotler, B.J., Hart, M.L., Long, D.G. & Forrest, L.L. 2016. Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate. New Phytologist 209(4):1734-46. doi: 10.1111/nph.13716

Söderström, L. –40 authors, J.C. Villarreal. 2016. World Checklist of hornworts and liverworts. Phytokeys 59: 1-828.


Chantanaorrapint, S., Penjor P & J.C. Villarreal. 2015. Taxonomic notes on Phaeoceros himalayensis, with the lectotypification of Anthoceros himalayensis. Phytotaxa 231(2): 193-196.

Forrest, L.L.,G. Long, P.M. Hollingsworth, D.C. Cargill & J.C. Villarreal. 2015. On Monocarpus (Monocarpaceae, Marchantiopsida), an isolated salt-pan complex thalloid liverwort allied to the Sphaerocarpales. Australian Systematic Botany 28: 137-144.

Li, F.-W., M. Melkonian, C.J, Rothfels, J.C. Villarreal, D.W. Stevenson, S.W. Graham, G. K-S. Wong, K.M. Pryer & S. Mathews. 2015. Phytochrome diversity in green plants and the origin of canonical plant phytochromes. Nature Communications 6: 7852 doi:10.1038/ncomms8852.

Villarreal, J.C. & K. Renzaglia. 2015. The hornworts: important advancements in early land plant evolution. Journal of Bryology 37(3): 157-170 (invited review).

For past publications by JCVA, click here.

Book Chapters and Tribute papers – peer reviewed

Hanson, D., K.S. Renzaglia & J.C. Villarreal. 2014. Diffusion limitation and CO2 concentrating mechanisms in bryophytes. In Advances in Photosynthesis and Respiration: Photosynthesis in Early Land Plants, D.T. Hanson & S.K. Rice (eds). Vol. 37: 95-112 Springer, Dordrecht. DOI:10.1007/978-94-007-6988-5_6.

Villarreal, J.C. 2013. Hornworts (genera Anthoceros and Phaeoceros). In The liverworts and hornworts of Tristan da Cunha, J. Váňa & J.J. Engel, J.J. (eds.). Memoirs of the New York Botanical Garden 105; 32-35, 85-86.

Renzaglia, K.S., J.C. Villarreal & R.J. Duff. 2009. New insights into morphology, anatomy and systematics of hornworts. In Bryophyte Biology II, B. Goffinet & J. Shaw (eds.), pp 139-171.

Edited volume

Villarreal, J.C., Frey & D.C. Cargill (eds.). 2010. Bryophyte Biology, Phylogeography, Systematics and Evolution in the Southern Hemisphere. Nova Hedwigia 91(3-4): 1–250 pp. (contributed peer-reviewed papers on biogeography, niche modeling, cryo-microscopy, floristics and evolution of Southern Hemisphere bryophytes).

Book reviews and Popular Publications

Villarreal, J.C. New insights on early land plant diversification: pyrenoid evolution and sexual system of hornworts. Field bryology 112: 55-57.

Pressel, S., J. Duckett & J.C. Villarreal. 2013. Hornwort heaven: recount of the expedition into the Himalayan foothills of northern Indian. Field bryology 110: 39-46.

Villarreal, J.C. & N. Salazar Allen. 2012. The enigmatic hornworts of the miniature forest of Cape Horn (English and Spanish). In Miniature forest of Cape Horn: Ecotourism with a hand lens. Goffinet, R. Rozzi, L. Lewis, W. Buck & F. Massardo (eds.) University of North Texas Press, pp. 118-126.

Villarreal, J.C. Before the vascular plants. Book review of “Syllabus of Plant Families A. Engler’s Syllabus der Pflanzenfamilien. 3. Bryophytes and seedless vascular plants.” 2009. Eds.: W. Frey, M. Stech & E. Fischer. The Bryologist 113: 431–434.

Posters in national and international meetings

Lavoie, C.*, et al. 2019. Une histoire de fidelité chez les lichens. Diversité et recrumente des symbiontes associés au lichen Stereocaulon spp. Colloque du Département de Biologie, Université Laval.

Conference Proceedings

Villarreal, J.C., Line Rochefort, Claire Boismenu & Melina Guêné-Nanchen. 2017. Future Arctic: A global intiative on bryophyte and lichen Arctic research: from species to ecosystems. May 24-26, 2017. Québec, Canada. Proceedings of the Workshop: Future Artic 2017