Leavitt, Steve


Email: 6yzk|kerkg|ozzFh{4kj{  (Email Form)

1115  MLBM
Provo, UT 84602
(801) 422-4879

Assistant Professor

 Teaching Interests
General biology, Symbioses, Evolution, Lichenology
 Research Interests
Evolutionary diversification, molecular systematics and phylogenomics, role of species interactions within an evolutionary context, fungal symbioses, biogeography in arid environments, conservation, climate change and bio-monitoring, and scientific education and outreach.
 Courses Taught

Winter 2013
  • BIO 100: Principles of Biology Section 004

Selected Publications

Journal Articles

Zhao X, Fernández-Brime S, Wedin M, Locke M, Leavitt SD, Lumbsch HT. 2017. Using multi-locus sequence data for addressing species boundaries in commonly accepted lichen-forming fungal species. 1–13. <website> doi:10.1007/s13127-016-0320-4

Leavitt SD, Divakar PK, Crespo A, Lumbsch HT. 2016. A matter of time – understanding the limits of the power of molecular data for delimiting species boundaries. Herzogia. 29(2):479– 492. <website>

Kirika PM, Divakar PK, Crespo A, Leavitt SD, Mugambi G, Gatheri GW, Lumbsch HT. 2016. Polyphyly of the genus Canoparmelia—uncovering incongruences between phenotype-based classification and molecular phylogeny within lichenized Ascomycota (Parmeliaceae). Phytotaxa. 289(1):036–048. <website> doi:http://dx.doi.org/10.11646/phytotaxa.289.1.2

Wei X, McCune B, Lumbsch HT, Li H, Leavitt SD, Yamamoto Y, Tchabanenko S, Wei J. 2016. Limitations of species delimitation based on phylogenetic analyses: a case study in the Hypogymnia hypotrypa group (Parmeliaceae, Ascomycota). 11(11):1-20. <website> doi:10.1371/journal.pone.0163664

Divakar PK, Leavitt SD, Molina MC, Del-Prado R, Lumbsch HT, Crespo A. 2016. A DNA barcoding approach for identification of hidden diversity in Parmeliaceae (Ascomycota): Parmelia sensu stricto as a case study. Botanical Journal of the Linnean Society. 180(1):21–29.  doi:10.1111/boj.12358

Alors D, Lumbsch HT, Divakar PK, Leavitt SD, Crespo A. 2016. An integrative approach for understanding diversity in the Punctelia rudecta species complex (Parmeliaceae, Ascomycota). PLoS ONE. 11(2):e0146537. <website> doi:10.1371/journal.pone.0146537

Lindgren H, Leavitt SD, Lumbsch HT. 2016. Characterization of microsatellite markers in the cosmopolitan lichen-forming fungus Rhizoplaca melanophthalma (Lecanoraceae). MycoKeys. 14:31-36. <website> doi:10.3897/mycokeys.14.9729

Leavitt SD, Kraichak E, Vondrak J, Nelsen MP, Sohrabi M, Perez-Ortega S, StClair LL, Lumbsch HT. 2016. Cryptic diversity and symbiont interactions in rock-posy lichens. Molecular Phylogenetics and Evolution. 99:261–274.  doi:10.1016/j.ympev.2016.03.030

Mark K, Saag L, Leavitt SD, Will-Wolf S, Nelsen MP, Tõrra T, Saag A, Randlane T, Lumbsch HT. 2016. Evaluation of traditionally circumscribed species in the lichen-forming genus Usnea, section Usnea (Parmeliaceae, Ascomycota) using a six-locus dataset. Organisms Diversity & Evolution. 16(3):497–524.  doi:10.1007/s13127-016-0273-7

Leavitt SD, Esslinger TL, Divakar PK, Crespo A, Lumbsch HT. 2016. Hidden diversity before our eyes: Delimiting and describing cryptic lichen-forming fungal species in camouflage lichens (Parmeliaceae, Ascomycota). Fungal Biology. 120(11):1374–1391.  doi:http://dx.doi.org/10.1016/j.funbio.2016.06.001

Kirika PM, Divakar PK, Crespo A, Gatheri GW, Mugambi G, Leavitt SD, Moncada B, Lumbsch HT. 2016. Molecular data show that Hypotrachyna sorocheila (Parmeliaceae) is not monophyletic. The Bryologist. 119(2):172–180. <website> doi:http://dx.doi.org/10.1639/0007-2745-119.2.172

Kirika PM, Divakar PK, Crespo A, Mugambi G, Orock EA, Leavitt SD, Gatheri GW, Lumbsch HT. 2016. Phylogenetic studies uncover a predominantly African lineage in a widely distributed lichen-forming fungal species. Mycokeys. 14:1–16.  doi:10.3897/mycokeys.14.8971

Widhelm TJ, Egan RS, Bertoletti FR, Asztalos MJ, Kraichak E, Leavitt SD, Lumbsch HT. 2016. Picking holes in traditional species delimitations: an integrative taxonomic reassessment of the Parmotrema perforatum group (Parmeliaceae, Ascomycota). Botanical Journal of the Linnean Society.  doi:10.1111/boj.12483

Leavitt SD, Grewe F, Widhelm TJ, Muggia L, Wray B, Lumbsch HT. 2016. Resolving evolutionary relationships in lichen-forming fungi using diverse phylogenomic datasets and analytical approaches. Scientific Reports. 6:22262.  doi:10.1038/srep22262

Lücking R, Hodkinson BP, Leavitt SD. 2016. The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota – Approaching one thousand genera. The Bryologist. 119(4):361-416.  doi:http://dx.doi.org/10.1639/0007-2745-119.4.361

Altermann S, Leavitt SD, Goward T. 2016. Tidying up the genus Letharia: introducing L. lupina sp. nov. and a new circumscription for L. columbiana. The Lichenologist. 48(5):423–439. <website> doi:http://dx.doi.org/10.1017/S0024282916000396

Zhao X, Leavitt SD, Zhao ZT, Zhang LL, Arup U, Grube M, Pérez-Ortega S, Printzen C, Sliwa L, Kraichak E, et al. 2016. Towards a revised generic classification of lecanoroid lichens (Lecanoraceae, Ascomycota) based on molecular, morphological and chemical evidence. Fungal Diversity. 78(1):293–304.  doi:10.1007/s13225-015-0354-5

Hawksworth DL, Lumbsch HT, Scholz P, Leavitt SD, Seaward MR. 2015. (2396) Proposal to conserve the name Lichen muralis (Lecanora muralis, Protoparmeliopsis muralis) with a conserved type (Ascomycota: Lecanorales: Lecanoraceae). Taxon. 64(6):1316–1317. <website> doi:https://doi.org/10.12705/646.18

Zhao X, Zhang LL, Zhao ZT, Wang WC, Leavitt SD, Lumbsch HT. 2015. A molecular phylogeny of the lichen genus Lecidella focusing on species from mainland China. PloS one. 10(9):e0139405. <website> doi:http://dx.doi.org/10.1371/journal.pone.0139405

Kraichak E, Divakar PK, Crespo A, Leavitt SD, Nelsen MP, Lücking R, Lumbsch HT. 2015. A tale of two hyper-diversities: diversification dynamics of the two largest families of lichenized fungi. Scientific Reports. 5:10028.  doi:10.1038/srep10028

Kantvilas G, Leavitt SD, Elix JA, Lumbsch HT. 2015. Additions to the genus Trapelia (Trapeliaceae: lichenised Ascomycetes). Australian Systematic Botany. 27(6):395–402.  doi:http://dx.doi.org/10.1071/SB14037

Singh G, Dal Grande F, Divakar PK, Otte J, Leavitt SD, Szczepanska K, Crespo A, Rico VJ, Aptroot A, da Silva Cáceres ME, et al. 2015. Coalescent-based species delimitation approach uncovers high cryptic diversity in the cosmopolitan lichen-forming fungal genus Protoparmelia (Lecanorales, Ascomycota). PloS one. 10(5):e0124625.  doi:http://dx.doi.org/10.1371/journal.pone.0124625

Divakar PK, Crespo A, Wedin M, Leavitt SD, Hawksworth DL, Myllys L, McCune B, Randlane T, Bjerke JW, Ohmura Y. 2015. Evolution of complex symbiotic relationships in a morphologically derived family of lichen-forming fungi. New Phytologist. 208(4):1217–1226.  doi:10.1111/nph.13553

Leavitt SD, Kraichak E, Nelsen MP, Altermann S, Divakar PK, Alors D, Esslinger TL, Crespo A, Lumbsch HT. 2015. Fungal specificity and selectivity for algae play a major role in determining lichen partnerships across diverse ecogeographic regions in the lichen-forming family Parmeliaceae (Ascomycota). Molecular Ecology. 24(14):3779–3797.  doi:10.1111/mec.13271

Buaruang K, Scharnagl K, Divakar PK, Leavitt SD, Crespo A, Manoch L, Lücking R, Lumbsch HT. 2015. Molecular data support Pseudoparmelia as a distinct lineage related to Relicina and Relicinopsis (Ascomycota, Lecanorales). The Lichenologist. 47(01):43–49. <website> doi:http://dx.doi.org/10.1017/S0024282914000577

Wang Q-, Yurkov AM, Gökerm M, Lumbsch HT, Leavitt SD, Groenewald M, Theelen B, Liu X-, Boekhout T, Bai F-. 2015. Phylogenetic classification of yeasts and related taxa within Pucciniomycotina. Studies in Mycology. 81:149–189.  doi:http://dx.doi.org/10.1016/j.simyco.2015.12.002

Kirika PM, Leavitt SD, Divakar PK, Crespo A, Gatheri GW, Mugambi G, Lumbsch HT. 2015. The monotypic genus Bulborrhizina belongs to Bulbothrix sensu lato (Parmeliaceae, Ascomycota). The Bryologist. 118(2):164–169. <website> doi:http://dx.doi.org/10.1639/0007-2745-118.2.164

Leavitt SD, Divakar PK, Ohmura Y, Wang L, Esslinger TL, Lumbsch HT. 2015. Who’s getting around? Assessing species diversity and phylogeography in the widely distributed lichen-forming fungal genus Montanelia (Parmeliaceae, Ascomycota). Molecular Phylogenetics and Evolution. 90:85–96. <website> doi:http://dx.doi.org/10.1016/j.ympev.2015.04.029

Leavitt SD, Esslinger TL, Hansen ES, Divakar PK, Crespo A, Loomis BF, Lumbsch HT. 2014. DNA barcoding of brown Parmeliae (Parmeliaceae) species: a molecular approach for accurate specimen identification, emphasizing species in Greenland. Organisms Diversity & Evolution. 14(1):11–20.  doi:10.1007/s13127-013-0147-1

Schoch CL, Robbertse B, Robert V, Vu D, Cardinali G, Irinyi L, Meyer W, Nilsson RH, Hughes K, Miller AN, et al. 2014. Finding needles in haystacks: linking scientific names, reference specimens and molecular data for Fungi. Database. 2014:bau061.  doi:10.1093/database/bau061

Altermann S, Leavitt SD, Goward T, Nelsen MP, Lumbsch HT. 2014. How do you solve a problem like Letharia? A new look at cryptic species in lichen-forming fungi using Bayesian clustering and SNPs from multilocus sequence data. PloS one. 9(5):e97556. <website> doi:http://dx.doi.org/10.1371/journal.pone.0097556

Shrestha G, Raphael J, Leavitt SD, StClair LL. 2014. In vitro evaluation of the antibacterial activity of extracts from 34 species of North American lichens. Pharmaceutical Biology. 52(10):1262–1266.  doi:10.3109/13880209.2014.889175

Esslinger TL, Hansen ES, Leavitt SD. 2014. The brown parmelioid lichen species in Greenland. Folia Cryptogamica Estonica. 51:25–48.  doi:http://dx.doi.org/10.12697/fce.2014.51.03

Nelsen MP, Thell A, Leavitt SD, Hampton-Miller CJ, Lumbsch HT. 2013. A reappraisal of Masonhalea (Parmeliaceae, Lecanorales) based on molecular and morphological data. The Lichenologist. 45(06):729–738.  doi:https://doi.org/10.1017/S0024282913000509

Leavitt SD, Lumbsch HT, StClair LL. 2013. Contrasting demographic histories of two species in the lichen-forming fungal genus Xanthomendoza (Teloschistaceae, Ascomycota). The Bryologist. 116(4):337–349.  doi:http://dx.doi.org/10.1639/0007-2745-116.4.337

Leavitt SD, Fernández-Mendoza F, Pérez-Ortega S, Divakar PK, Lumbsch HT, StClair LL. 2013. DNA barcode identification of lichen-forming fungal species in the Rhizoplaca melanophthalma species-complex (Lecanorales, Lecanoraceae), including five new species. MycoKeys. 7:1.  doi:https://doi.org/10.3897/mycokeys.7.4508

Leavitt SD, Esslinger TL, Nelsen MP, Lumbsch HT. 2013. Further species diversity in Neotropical Oropogon (Lecanoromycetes: Parmeliaceae) in Central America. The Lichenologist. 45(04):553–564.  doi:https://doi.org/10.1017/S0024282913000212

Parnmen S, Leavitt SD, Rangsiruji A, Lumbsch HT. 2013. Identification of species in the Cladia aggregata group using DNA barcoding (Ascomycota: Lecanorales). Phytotaxa. 115(1):1–14.  doi:http://dx.doi.org/10.11646/phytotaxa.115.1.1

Leavitt SD, Fernández-Mendoza F, Pérez-Ortega S, Sohrabi M, Divakar PK, Vondrák J, Lumbsch HT, StClair LL. 2013. Local representation of global diversity in a cosmopolitan lichen-forming fungal species complex (Rhizoplaca, Ascomycota). Journal of Biogeography. 40(9):1792–1806.  doi:10.1111/jbi.12118

Leavitt SD, Esslinger TL, Spribille T, Divakar PK, Lumbsch HT. 2013. Multilocus phylogeny of the lichen-forming fungal genus Melanohalea (Parmeliaceae, Ascomycota): Insights on diversity, distributions, and a comparison of species tree and concatenated topologies. Molecular Phylogenetics and Evolution. 66(1):138–152.  doi:http://dx.doi.org/10.1016/j.ympev.2012.09.013

Leavitt SD, Lumbsch HT, Stenroos S, StClair LL. 2013. Pleistocene speciation in North American lichenized fungi and the impact of alternative species circumscriptions and rates of molecular evolution on divergence estimates. PloS one. 8(12):e85240.  doi:http://dx.doi.org/10.1371/journal.pone.0085240

Leavitt SD, Nelsen MP, Lumbsch HT, Johnson LA, StClair LL. 2013. Symbiont flexibility in subalpine rock shield lichen communities in the Southwestern USA. The Bryologist. 116(2):149–161.  doi:http://dx.doi.org/10.1639/0007-2745-116.2.149

Sohrabi M, Leavitt SD, Rico VJ, Halici MG, Shrestha G, Stenroos S. 2013. Teuvoa, a new lichen genus in Megasporaceae (Ascomycota: Pertusariales), including Teuvoa junipericola sp. nov. 45(03):347–360.  doi:https://doi.org/10.1017/S0024282913000108

Leavitt SD, Nelsen MP, StClair LL. 2013. Treading in murky waters: Making sense of diversity in Xanthoparmelia (Parmeliaceae, Ascomycota) in theWestern United States. Bulletin of the California Lichen Society. 19(2):58–69.

Divakar PK, Kauff F, Crespo A, Leavitt SD, Lumbsch HT. 2013. Understanding phenotypical character evolution in parmelioid lichenized fungi (Parmeliaceae, Ascomycota). PloS one. 8(11):e83115.  doi:http://dx.doi.org/10.1371/journal.pone.0083115

Shrestha G, Leavitt SD, Proulx MW, Glacy LA, Call C, Hendrickson J, StClair LL. 2012. A Checklist of the lichens of the Beaver Dam Slope, Washington County, Utah, USA. North American Fungi. 7:1–7.  doi:http://dx.doi.org/10.2509/naf2012.007.005

Esslinger TL, Morse CA, Leavitt SD. 2012. A new North American species of Hyperphyscia (Physciaceae). The Bryologist. 115(1):31–41.  doi:http://dx.doi.org/10.1639/007-2745-115.1.31

Thell A, Crespo A, Divakar PK, Kärnefelt I, Leavitt SD, Lumbsch HT, Seaward MR. 2012. A review of the lichen family Parmeliaceae–history, phylogeny and current taxonomy. Nordic Journal of Botany. 30(6):641–664.  doi:10.1111/j.1756-1051.2012.00008.x

Divakar PK, Del-Prado R, Lumbsch HT, Wedin M, Esslinger TL, Leavitt SD, Crespo A. 2012. Diversification of the newly recognized lichen-forming fungal lineage Montanelia (Parmeliaceae, Ascomycota) and its relation to key geological and climatic events. American Journal of Botany. 99(12):2014–2026.  doi:10.3732/ajb.1200258

Orock EA, Leavitt SD, Fonge BA, StClair LL, Lumbsch HT. 2012. DNA-based identification of lichen-forming fungi: can publicly available sequence databases aid in lichen diversity inventories of Mount Cameroon (West Africa)?. The Lichenologist. 44(06):833–839.  doi:https://doi.org/10.1017/S0024282912000424

Leavitt SD, Esslinger TL, Divakar PK, Lumbsch HT. 2012. Miocene and Pliocene dominated diversification of the lichen-forming fungal genus Melanohalea (Parmeliaceae, Ascomycota) and Pleistocene population expansions. BMC Evolutionary Biology. 12(1):1.  doi:10.1186/1471-2148-12-176

Leavitt SD, Esslinger TL, Lumbsch HT. 2012. Neogene-dominated diversification in neotropical montane lichens: dating divergence events in the lichen-forming fungal genus Oropogon (Parmeliaceae). American Journal of Botany. 99(11):1764–1777.  doi:10.3732/ajb.1200146

Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA, Chen W, Bolchacova E, Voigt K, Crous PW, et al. 2012. Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proceedings of the National Academy of Sciences. 109(16):6241–6246.  doi:10.1073/pnas.1117018109

Leavitt SD, Johnson L, StClair LL. 2011. Species delimitation and evolution in morphologically and chemically diverse communities of the lichen-forming genus Xanthoparmelia (Parmeliaceae, Ascomycota) in western North America. American Journal of Botany. 98(2):175–188.

Leavitt SD, StClair LL. 2008. Lichens of the Boulder Mountain Plateau, Wayne County, Utah, USA. Evansia. 25(4):85–89.

Jackson HB, Leavitt SD, Krebs T, StClair LL. 2005. Lichen flora of the eastern Mojave Desert: Blackrock Arizona, Mojave County, Arizona, USA. Evansia. 22(1):30–38.


Leavitt SD, Lumbsch HT. 2016. Ecological Biogeography of Lichen-Forming Fungi.  doi:10.1007/978-3-319-29532-9_2

Leavitt SD, StClair LL. 2015. Bio-monitoring in Western North America: What Can Lichens Tell Us About Ecological Disturbances?.  doi:10.1007/978-81-322-2181-4_5

Leavitt SD, Moreau CS, Lumbsch HT. 2015. The dynamic discipline of species delimitation: progress toward effectively recognizing species boundaries in natural populations.  doi:10.1007/978-81-322-2235-4_2

Divakar P, Lumbsch H, Wedin M, Bjerke J, McCune B, Kauff F, Roca-Valiente B, Nunes-Zapata J, del Prado R, Ruibal C. 2012. Multilocus phylogeny and classification of Parmeliaceae (Ascomycota) derived from Parsys-10.

Technical Report

StClair LL, Leavitt SD. 2013. Final Report Concerning Establishment of a Lichen Air Quality Bio-monitoring Program for the Quinn Canyon and Grant Range Wilderness Areas, Humboldt Toiyabe National Forest, Nevada.

StClair LL, Leavitt SD. 2013. Final Report Concerning Establishment of a Lichen Air Quality Bio-monitoring Program for the South Unit of the Duchesne Ranger District, Ashley National Forest and Detailed Elemental Analysis Survey of the Cart Creek Bridge and Selected Sites along U.S. Highway 191, Utah.

StClair LL, St Clair SB, Leavitt SD. 2013. Final Report concerning Review of the lichen air quality baseline for Region 1 of the Forest Service, Montana and Idaho.

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