Northwest Lichenologists

Recent papers revising macrolichens in the Pacific Northwest

The following papers were published since McCune and Geiser's (2009) second edition of Macrolichens of the Pacific Northwest. This list contains revisions of species concepts, major range extensions (e.g., new reports for North America based on specimens from the Pacific NW, and new keys. I have probably forgotten some items, so please email corrections to me.

Revisions of crustose lichens are too numerous to list. Please see the world list of recent lichen literature and the Guide to Literature.

Authors! Please note how some journals are freely available to all, specifically Graphis Scripa, MycosphereNorth American Fungi, and Opuscula Philolichenum. Please support the free, global distribution of scientific information by publishing in these journals.

Altermann, S., S. D. Leavitt & T. Goward. 2016. Tidying up the genus Letharia: introducing L. lupina sp. nov. and a new circumscription for L. columbiana. Lichenologist 48:423-440.

Arcadia, L. 2013. Usnea dasopoga, a name to be reinstated for U. filipendula, and its orthography. Taxon 62:604-605. This change is strictly nomenclatural and does not address species concepts. Because both the original authors' intentions and the International Botanical Code of Nomenclature are open to interpretation, do not be surprised if this name changes again.

Athukorala, S. N. P. et al. 2016. Phylogenetic relationships among reindeer lichens of North America. Lichenologist 48:209-228.  Some traditional species concepts are supported, others are not (e.g. Cladonia stygia vs. C. rangiferina and C. arbuscula vs. C. mitis

Bjork, C. R. 2009. Additions to the lichen flora of Washington state, United States. I. Evansia 27:18-20. Macrolichens added were Leptogium schraderi, Massalongia microphylliza, Physconia leucoleiptes, and Solorina bispora. Crustose species were added as well.

Boluda, C. G., et al. 2016. Microchemical and molecular investigations reveal Pseudephebe species as cryptic with an environmentally modified morphology. Lichenologist 48:527-544.

Brodo, I. M., C. Freebury & N. Alfonso. 2013. Notes on the lichens Physcia aipolia and Physcia alnophila in North America. Evansia 30: 110-119. Practical advice on using morphological and chemical characters to separate the recently recognized P. alnophila from P. aipolia. The dominant taxon in the PNW is P. alnophila.

Carlsen, T. & 7 other authors. 2012. Species delimitation, bioclimatic range, and conservation status of the threatened lichen Fuscopannaria confusa. Lichenologist 44: 565-575.  (Although the data in this paper are Scandinavian, the results might apply to our records of these species as well. F. confusa, F. ahlneri, F. mediterranea, and Parmeliella parvula are shown to be distinct, but F. confusa and P. parvula cannot be distinguished in their normally sterile condition.)

Davydov, E. A., D. E. Himelbrant & I. S. Stepanchikova. 2011. Contribution to the study of Umbilicariaceae (lichenized Ascomycota) in Russia. II. Kamchatka Peninsula. Herzogia 24: 251–263. Umbilicaria krascheninnikovii is Umbilicaria polaris for N Am records.

Divakar, P. K.,R. Del-Prado, H. T. Lumbsch, M. Wedin, T. L. Esslinger, S. D. Leavitt, & A. Crespo. 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 99:2014–2026. (The new genus Montanelia is erected to include the saxicolous species Melanelia disjuncta, M. panniformis, M. sorediata, & M. tominii, among others.

Divakar, P. K., G. Figueras, N. L. Hladun & A. Crespo. 2010. Molecular phylogenetic studies reveal an undescribed species within the North American concept of Melanelixia glabra (Parmeliaceae). Fungal Diversity 42: 47-55.  (North American material is shifted to M. californica A. Crespo & Divakar.)

Divakar, P. K., A. Crespo, E. Kraichak, S. D. Leavitt, G. Singh, I. Schmitt, & H. T. Lumbsch. 2017. Using a temporal phylogenetic method to harmonize family and genus-level classification in the largest clade of lichen-forming fungi. Fungal Diversity 84:101. DOI 10.1007/s13225-017-0379-z  (The numerous generic splits in Cetraria have long been problematic; this paper fixes those problems and simplifies the classification. Most cetrarioid lichens are assigned to either Nephromopsis or Cetraria. The following names (among others) are reduced to synonymy: Ahtiana, Allocetraria, Flavocetraria, Kaernefeltia, Masonhalea, Tuckermanella, Tuckermannopsis, and Vulpicida. Gowardia is reunited with Alectoria. One problem with the new classification proposed by Divakar et al. is that Cetraria subalpina was apparently misplaced into Nephromopsis. Based on phylogenetic trees by Thell, Cetraria subalpina should remain in Cetraria. See pdf or Excel spreadsheet of changes relevant to the Pacific Northwest.)

Dolnik, C., A. Beck, & D. Zarabska. 2010. Distinction of Cladonia rei and C. subulata based on molecular, chemical and morphological characteristics. Lichenologist 42: 373-386.  (These two species are recognized as distinct, in contrast to Spier & Aptroot (2008).)

Esslinger, T. L. & K. L. Dillman. 2010. Physconia grumosa in North America. Bryologist 113:77-80. (This Asian species is now reported from Alaska, central and southern Rocky Mts, and eastern states and provinces).

Glavich, D. 2009. Distribution, rarity and habitats of three aquatic lichens on federal land in the U.S. Pacific Northwest. Bryologist 112:54-72. (Peltigera hydrothyria, Leptogium rivale, Dermatocarpon meiophyllizum )

Goward, T., T. Ahti, J. A. Elix & T. Spribille. 2010. Hypogymnia recurva and Hypogymnia wilfiana spp. nov., two new lichens from western North America. Botany 88:345-361. (North American material is separated as H. wilfiana from the Japanese species H. metaphysodes.)

Goward, T., T. Spribille, T. Ahti & C. J. Hampton-Miller. 2012. Four new sorediate species in the Hypogymnia austerodes group (lichens) from northwestern North America, with notes on thallus morphology. Bryologist 115:84-100. (H. dichroma, H. protea, H. salsa, and H. verruculosa spp. nov.)

Halonen, P., L. Myllys, S. Velmala, & H. Hyvarinen. 2009. Gowardia (Parmeliaceae)-- a new alectorioid lichen genus with two species. Bryologist 112:138-146.

Hodkinson, B. P., J. C. Lendemer, & T. L. Esslinger. 2010. Parmelia barrenoae a macrolichen new to North America and Africa. North American Fungi 5(3):1-5. (This species is a segregate of Parmelia sulcata, distinguishable both morphologically and by ITS sequence.)

Hodkinson, B. P. & J. C. Lendemer. 2011. Molecular analyses reveal semi-cryptic species in Xanthoparmelia tasmanica. Bibliotheca Lichenologica 106:115-126. (North American material is referred to X. hypofusca (Gyeln.) Hodkinson & Lendemer, X. tasmanica is strictly austral)

Jørgensen, P. M. 2008. Vahliella, a new lichen genus. Lichenologist 40:221-225.

New combinations relevant to PNW:
  • Vahliella californica (Tuck.) P. M. Jørg.
  • Vahliella hookerioides (P. M. Jørg.) P. M. Jørg.
  • Vahliella leucophaea (Vahl) P. M. Jørg.
  • Vahliella saubinetii (Mont.) P. M. Jørg.

Jørgensen, P. M. & T.Goward. 2015. Notes on the Collema tenax complex, with recognition of two further species in the genus Enchylium (Ach.) Gray. Evansia 32: 42-47. Enchylium expansum (Degel.) P. M. Jørg. (spreading thallus, 3-septate spores, sometimes isidiate; moist calcareous rock, arctic-alpine). Enchylium substellatum (H. Magn.) P. M. Jørg. (cordilleran, calciphile, 3-septate spores, longer and narrower than E. tenax.

Jørgensen, P. M. & T. Tonsberg. 2010. Leptogium insigne, a new species from the Pacific Northwest of North America. Bibliotheca Lichenologica 104:241-245. (this new species is segregated from L. brebissonii, a subtropical species that does not occur in the PNW.)

Knudsen, K. & J. C. Lendemer. 2009. Cladonia maritima, a new species in the C. cervicornis group from western North America. Opuscula Philolichenum 6:121-124. (segregate species from southern California; includes key to C. cervicornis group in California.)

Leavitt, S. D., T. L. Esslinger, P. K. Divakar & H. T. Lumbsch. 2012. Miocene divergence, phenotypically cryptic lineages, and contrasting distribution patterns in common lichen-forming fungi (Ascomycota: Parmeliaceae). Biological Journal of the Linnean Society 107: 920-937. (Most specimens in western North America have been misplaced in M. fuliginosa (Fr. ex Duby) O. Blanco et al. M. glabratula is commonly corticolous, but also occurs on rock. M. glabratula appears to be the only one of the two taxa that occurs in North America.)

Leavitt, S. D., et al. 2013. DNA barcode identification of lichen-forming fungal species in the Rhizoplaca melanopthalma species-complex (Lecanoralies, Lecanoraceae), including five new species. MycoKeys 7:1-22. Describes new species recognizable by DNA sequences but with no morphological distinctions.

Leavitt, S. D., P.K. Divakar, A. Crespo & H. Thorsten Lumbsch. 2016. A matter of time – understanding the limits of the power of molecular data for delimiting species boundaries. Herzogia 29:479 – 492.  (Discussion and re-analysis of species-level results in Bryoria and Thamnolia).

Lendemer, J. C. & B. P. Hodkinson. 2010. A new perspective on Punctelia subrudecta (Parmeliaceae) in North America: previously rejected morphological characters corroborate molecular phylogenetic evidence and provide insight into an old problem. Lichenologist 42: 405-421.  (Material of the P. subrudecta group in the PNW is refered to P jeckeri (Roum.) Kalb; eastern North America has several species in this group.)

Lendemer, J. C. & H. O'Brien. 2011. How do you reconcile molecular and non-molecular datasets? A case study where new molecular data prompts a revision of Peltigera hydrothyria s.l. in North America and the recognition of two species. Opuscula Philolichenum 9:99-110. (Western specimens are placed in P. gowardii sp. nov.)

Lumbsch, H. T. & many other authors. 2011. One hundred new species of lichenized fungi: a signature of undiscovered global diversity. Phytotaxa 18: 1-127. Hypogymnia amplexa Goward, Björk & Wheeler, sp. nov. and Platismatia wheeleri Goward, Altermann & Björk, sp. nov.

Magain, N., E. Sérusiaux, M. P. Zhurbenko, F. Lutzoni & J. Miadlikowska. 2016a. Disentangling the Peltigera polydactylon species complex by recognizing two new taxa, P. polydactylon subsp. udeghe and P. seneca. Herzogia 29:514-528. (New subspecies that is widespread in western North America.)

Marcelli, M. P., L. S. Canez, M. N. Benatti, A. A. Spielmann, P. Jungbluth & J. A. Elix. 2011. Taxonomic novelties in Parmeliaceae. Bibliotheca Lichenologica 106:115-126. Parmotrema herrei (Zahlbr.) Spielmann & Marcelli is known from only from the type locality (collected in 1906 and 1942), on sandstone crevices in the Santa Cruz Mountains in coastal California. It has not been relocated, but should be sought elsewhere on the Pacific Coast. "...recognized by the long, thickened and branched cilia distributed along the lacinules, together with the strictly laciniate thallus," absence of soredia and isidia, and presence of salacinic acid.

Mark, K. et al. (8 authors). 2016. Evaluation of traditionally circumscribed species in the lichen-forming genus Usnea, section Usnea (Parmeliaceae, Ascomycota) using a six-locus dataset. Organisms Diversity & Evolution (no volume number) 1-28. Usnea substerilis is synonymized under U. lapponica. Some traditional morphospecies are supported, e.g. U. cavernosa, U. silesiaca, U. wasmuthii.

Miadlikowska, J., C. L. Schoch, S. A. Kageyama, K. Molnar, F. Lutzoni & B. McCune. 2011. Hypogymnia phylogeny, including Cavernularia, reveals biogeographic structure. The Bryologist 114:392-400. doi:10.1639/0007-2745-114.2.392. (Our two Cavernulia species are shown to belong to Hypogymnia, H. lophyrea (Ach.) Krog and H. hultenii (Degel.) Krog as proposed by Hildur Krog in 1951.)

Miadlikowska J., Richardson D., Magain N., Ball B., Anderson F., Cameron R., Lendemer J., Truong C., Lutzoni F. 2014. Phylogenetic placement, species delimitation, and cyanobiont identity of endangered aquatic Peltigera species (lichen-forming Ascomycota, Lecanoromycetes). American Journal of Botany 101:1141-1156.  What used to be considered a single species, Hydrothyria venosa, then two species, Peltigera gowardii (western N Am) and Peltigera hydrothyria (eastern N Am), is confirmed to have three supported clades. The third species is described formallly, Peltigera aquatica, an action that is sure to be controversial among practicing field lichenologists in the PNW, simply because P. aquatica occurs with P. gowardii, but is indistinguishable morphologically or chemically. This action has made it impossible to identify western Hydrothyria without DNA sequencing, if you accept the three-species documented by this paper.

McCune, B., J. Di Meglio & M. J. Curtis. 2014. An unusual ascospore shape and a new species, Umbilicaria nodulospora (Umbilicariaceae), from California and Oregon. Bryologist 117:170-178.

McCune, B., R. Rosentreter, T. Spribille, O. Breuss & T. B. Wheeler. 2014. Montana Lichens: An Annotated List. Monographs in North American Lichenology 2:1-183. Northwest Lichenologists, Corvallis, Oregon. New combinations: Lobaria anomala (Brodo & Ahti) Spribille & McCune, Lobaria anthraspis (Ach.) Spribille & McCune, Rostania quadrifida (Stone & McCune) McCune, Scytinium tacomae (P. M. Joerg. & Toensb.) McCune.

McCune, B., C. Schoch, H. T. Root, S. A. Kageyama & J. Miadlikowska. 2011. Geographic, climatic, and chemical differentiation in the Hypogymnia imshaugii species complex (Parmeliaceae). Bryologist 114:526-544. ("Segregation of the morphotype described as H. amplexa is not supported by our chemotype or ITS groupings." H. amplexa is considered a synonym of H. imshaugii.)

Miller, J. E. D. 2011. The Usnea rigida group in California and the Pacific Northwest. Bulletin of the California Lichen Society 18:1-4. (Discusses the origin of differences in recent taxonomic concepts of Usnea rigida and relatives, including U. arizonica, U. florida, U. intermedia, and U. quasirigida.)

Moberg, R. 2011. The lichen genus Heterodermia (Physciaceae) in South America -- a contribution including five new species. Nordic Journal of Botany 29:129-147. Contains full descriptions and many species, including the rare (with us) Heterodermia japonica.

Molina, M. C., P. K. Divakar, T. Goward, A. M. Millanes, H. T. Lumbsch & A. Crespo. 2017. Neogene diversification in the temperate lichen-forming fungal genus Parmelia (Parmeliaceae, Ascomycota). Systematics and Biodiversity 15:166-181. (New: Parmelia sulymae, primarily epiphytic, similar to P. hygrophila, and P. imbricaria, primarily saxicolous.)

Moncada, B., R. Lucking & L. Betancourt-Macuase. 2013. Phylogeny of the Lobariaceae (lichenized Ascomycota: Peltigerales) with a reappraisal of the genus Lobariella. Lichenologist 45:203-263. The authors take a narrow view of Lobaria, recognizing several segregate genera, rather than adopting a broad view of the genus. New combinations: Anomalobaria anomala, A. anthraspis, Dendriscosticta oroborealis, Dendriscosticta wrightii. Also recognized are Lobarina scrobiculata and Lobarina oregana. Other related Pacific Northwest species (Lobaria hallii, Pseudocyphellaria rainierensis) were not treated because existing DNA sequence data were deemed unreliable.

Moncada,, B., B. Reidy & R. Lucking. 2014. A phylogenetic revision of Hawaiian Pseudocyphellaria sensu lato (lichenized Ascomycota: lobariaceae) reveals eight new species and a high degree of inferred endemism. Bryologist 117:119-160. (Pseudocyphellaria perpetua is synonymized with P. hawaiiensis.)

Muggia, L., P. Nelson, T. Wheeler, L. S. Yakovchenko, T. Tonsberg & T. Spribille. 2011. Convergent evolution of a symbiotic duet: the case of the lichen genus Polychidium (Peltigerales, Ascomycota). American Journal of Botany 98:1647-1656. (The lichen that we have called Polychidium contortum in the PNW is, according to these authors, Leptogidium dendriscum. Leptogidium has jigsaw puzzle-like cortex while Polychidium has roundish cells in the cortex. The type of L. dendriscum is, however, from Tahiti, so it is quite unlikely that this is the same species as in the PNW. Meanwhile, they have applied L. contortum to Austral material (New Zealand, Chile) and L. dendriscum to northern material. This should be considered provisional until material is studied from throughout the range of the genus.)

Myllys, L., P. Halonen & S. Velmala. 2006. Notes on some rare species of Bryoria from Finland. Graphis Scripta 18:23-26. (Bryoria americana (Motyka) Holien is recognized at the species level following Holien. Also discussed are B. glabra, B. tortuosa, and B. tenuis. Continued recognition of B. americana at the subspecies level by McCune & Geiser (2009) as B. trichodes ssp. americana was an oversight rather than a conscious decision.)

Myllys, L., S. Velmala, H. Holien, P. Halonen, L.-S. Wang & T. Goward. 2011. Phylogeny of the genus Bryoria. Lichenologist 43:617-638.

Myllys, L. et al. 2015. Taxonomic delimitation of the genera Bryoria and Sulcarea, with a new combination Sulcaria spiralifera introduced. Lichenologist 46:737-752. Bryoria spiralifera moved to Sulcaria and B. pseudocapillaris is synonymized with S. spiralifera.

Myllys, L. et al. 2016. New species in Bryoria (Parmeliaceae, Lecanoromycetes) from north-west North America. Lichenologist 48:355-366. New: Bryoria alaskana, B. irwinii

Myllys, L. et al. 2016.Chemical diversity and ecology of the genus Bryoria section Implexae (Parmeliaceae) in Finland. Bryologist 119:29-38. "...chemotypes in the section have distinct ecological and climatic preferences even if they lack genetic variation as previously shown by Velmala et al. (2014)."

Nelsen, M. P., N. Chavez, E. Sackett-Hermann, A. Thell, T. Randlane, P. K. Divakar, V. J. Rico and H. T. Lumbsch. 2011. The cetrarioid core group revisited (Lecanorales: Parmeliaceae). Lichenologist 43: 537-551. (Cetrarioid genera are still in disarray: "The present study largely confirms the topologies of previous studies, but with increased support. Approximately half of the genera in the cetrarioid core were not monophyletic.")

Nelsen, M. P. & A. Gargas. 2009. Assessing clonality and chemotype monophyly in Thamnolia (Pertusariales: Icmadophilaceae). Bryologist 112:42-53. (Concludes that chemotypes in T. vermicularis do not form strongly supported monophyletic groups, thus T. subuliformis is considered conspecific with T. vermicularis.)

Nelson, P., J. Walton & C. Roland. 2009. Erioderma pedicellatum (Hue) P. M. Jorg, new to the United States and western North America, discovered in Denali National Park and Preserve and Denali State Park, Alaska. Evansia 26:19-21.

Nelson, P., J. Walton, H. Root & T. Spribille. 2011. Hypogymnia pulverata(Parmeliaceae) and Collema leptaleum (Collemataceae), two macrolichens new to Alaska. North American Fungi 6(7):1-8.

Onut-Brannstrom, I., L. Tibell & H. Johannesson. 2017. A worldwide phylogeography of the whiteworm lichens Thamnolia reveals three lineages with distinct habitats and evolutionary histories. Ecology and Evolution 7:3602-3615. ("... we did not find support for the recognition of taxa in Thamnolia based on either morphological or chemical characters." Compare wtih Leavitt et al. (2016).)

Otalora, M. A. G., I. Martinez, M. Carmen Molina, G. Aragon & F. Lutzoni. 2008. Phylogenetic relationships and taxonomy of the Leptogium lichenoides group (Collemataceae, Ascomycota) in Europe. Taxon 57:907-921. (Much of the material from the Pacific NW appears to fit their concept of L. pulvinatum.)

Pino-Bodas, R., A. R. Burgaz, M. P. Martin & H. T. Lumbsch. 2011. Phenotypical plasticity and homoplasy complicate species delimitation in the Cladonia gracilis group (Cladoniaceae, Ascomycota). Organisms Diversity and Evolution 11:343-355. (The distinction between C. ochrochlora and C. coniocraea is not supported. C. coniocraea is the older name and thus C. ochrochlora can be treated as a synonym of C. coniocraea.)

Pino-Bodas, R., A. R. Burgaz, M. P. Martin & H. T. Lumbsch. 2012. Species delimitation in the Cladonia cariosa group (Cladoniaceae, Ascomycota). Lichenologist 44:121-135. (Cladonia dahliana is not monophyletic, according to phylogenetic reconstructions based on three loci. The taxon is more appropriately placed as the psoromic acid chemotype of C. symphycarpa.)

Pino-Bodas, R., et al. 2015. The phenotypic features used for distinguishing species within the Cladonia furcata complex are highly homoplasious. Lichenologist 47:287-303. Cladonia furcata is polyphyletic. The characters usually used to distinguish species in this group to not accord well with phylogenetic reconstructions. The one specimen of C. furcata from the PNW that was sequenced does not appear in the trees. C. scabriuscula and C. furcata do not separate in the trees. The study stopped short of making taxonomic revisions.

Spribille, T. et al. Contributions to an epiphytic lichen flora of northwest North America: I. Eight new species from British Columbia inland rain forests. Bryologist 112:109-137. (Collema coniophilum Goward, very similar to C. furfuraceum, described from two areas in eastern BC).

Stone, D. & B. McCune. 2010. Collema quadrifidum, a new epiphytic lichen species from the Pacific Northwest of the United States. North American Fungi 5(2):1-6.  doi: 10.2509/naf2010.005.002

Stone, D. F., J. W. Hinds, F. L. Anderson & J. C. Lendemer. 2016. A revision of the Leptogium saturninum group in North America. Lichenologist 48:387-422. Thought you had L. saturninum? Better check again! New PNW species: L. compactum, L. cookii.

Thell, A., F. Hognabba, J. A. Elix, T. Feuerer, I. Karnefelt, L. Myllys, T. Randlane, A. Saag, S. Stenroos, T. Ahti & M. R. D. Seaward. 2009. Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers. Lichenologist 41:489-512. (This paper clearly shows the taxonomic chaos still present in Cetraria in the broad sense. Most of the Cetrarioid genera are not supported by analysis of genetic data. See also Nelsen et al. (2011))

Timdal, E. & T. Tonsberg. 2012. Cladonia straminea, the correct name for C. metacorallifera. Graphis Scripta 24:33-35.

Velmala, S., L. Myllys, P. Halonen, T. Goward & T. Ahti. 2009. Molecular data show that Bryoria fremontii and B. tortuosa (Parmeliaceae) are conspecific. Lichenologist 41:231-242.

Velmala et al. 2014. Taxonomy of Bryoria section Implexae (Parmeliaceae, Lecanoromycetes) in North America and Europe, based on chemical, morphological and molecular data. Ann. Bot. Fennici 51:345-371. The genetic data show that North American Bryoria pikei, pseudofusecens, and friabilis all belong to the same species, showing almost no genetic variation with a 3-locus data set, yet a new species, B. inactiva, is described within that single clade. A broader view that would be supported by the data (but not taken by the authors) is that all four belong to a single species, and that the chemotypes might be given taxonomic rank below the species level (e.g. varieties).

Vondrak, J. et al. 2013. Two superficially similar lichen crusts, Gregorella humida and Moelleropsis nebulosa, and a description of the new lichenicolous fungus Llimoniella gregorellae. Herzogia 26:31-48. Moelleropsis nebulosa (now in Fuscopannaria) is hardly a macrolichen but it is included in McCune & Geiser (2009).

Westberg, M. & U. Arup. 2011. Candelaria pacifica sp. nova (Ascomycota, Candelariales) and the identity of Candelaria vulgaris. Bibliotheca Lichenologica 106:353-364. (Validates the name C. pacifica, which is the common species of Candelaria in the Pacific coastal states, but ranging widely in western North America. Candelaria pacifica discussed and illustrated in McCune & Geiser (2009), but at that time the species had not been formally described.)

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