Bulletin of the California Lichen Society

Volume 1 9 No. 2 Winter 201 2 The California Lichen Society seeks to promote the appreciation, conservation, and study of lichens. The interests of the Society include the entire western part of the continent, although the focus is on California. Dues categories are (in $US per year): Student and fixed income (eBulletin only) ­ $10, Regular ­ $20 ($25 for foreign members, or $20 foreign student memberships), Family ­ $25, Sponsor and Libraries ­ $35, Donor ­ $50, Benefactor ­ $100, and Life Members ­ $500 (one time) payable to the California Lichen Society, PO Box 472, Fairfax, California 94978. Members receive the Bulletin and notices of meetings, field trips, lectures and workshops.

Board Members of the California Lichen Society: President: Bill Hill, [email protected] Vice President: Shelly Benson, [email protected] Secretary: Erin P. Martin, [email protected] Treasurer: Kathy Faircloth, [email protected] Member At Large: Tom Carlberg, [email protected]

Editor: John Villella, [email protected] Production Editor: Erin P. Martin, [email protected]

Committees of the California Lichen Society: Conservation: Eric Peterson, [email protected], chairperson Education: Erin P. Martin, [email protected], chairperson Poster/Mini Guides: Susan Crocker, [email protected], chairperson Events/Field trips/Workshops: vacant, chairperson Outreach: vacant, chairperson

The Bulletin of the California Lichen Society (ISSN 1093­9148) is edited by John Villella ([email protected]) and is produced by Erin P. Martin ([email protected]). The Bulletin welcomes manuscripts on technical topics in lichenology relating to western North America and on conservation of the lichens, as well as news of lichenologists and their activities. The best way to submit manuscripts is by e­mail attachments in the format of a major word processor (DOC or RTF preferred). Do include italics for scientific names. Please submit figures in electronic formats with a resolution of 300 pixels per inch (600 minimum for line drawings). Email submissions are limited to 10MB per email, but large files may be split across several emails or other arrangements can be made. Contact the Production Editor: Erin P. Martin, at [email protected], for details of submitting illustrations or other large files. A review process is followed. Nomenclature follows Esslingers cumulative checklist online at http://www.ndsu.nodak.edu/instruct/essinge/chchlist/chcklist7.htm. The editors may substitute abbreviations of authors names, as appropriate from R.K. Brummitt and C.E. Powell, Authors of Plant Names, Royal Botanic Gardens, Kew, 1992. Style follows this issue. Electronic reprints in PDF format will be emailed to the lead author at no cost.

The deadline for submitting material for the Summer 2013 CALS Bulletin is 31 March 2013.

The California Lichen Society is online at: http://californialichens.org/ and has email discussions through http://tech.groups.yahoo.com/group/CaliforniaLichens/.

Volume 19 (2) of the Bulletin was issued on 15 December 2012.

Front Cover: Xanthoparmelia aff. chlorochroa with Sedum lanceolatum and Phlox pulvinata, Aquarius Plateau, Utah, USA. See article by Leavitt et al. pg 1. Photo by Steven Leavitt. Bulletin of the California Lichen Society

VOLUME 1 9 NO. 2 Winter 201 2

Treading in murky waters: Making sense of diversity in Xanthoparmelia (Parmeliaceae, Ascomycota) in the Western United States

Steven D. Leavitt1, Matthew P. Nelsen2, and Larry L. St. Clair3 1The Field Museum, Department of Botany, 1400 South Lake Shore Drive, Chicago, Illinois 60605 ­ [email protected] 2University of Chicago, Committee on Evolutionary Biology, 1025 E. 57th Street, Chicago, Illinois 60637 ­ [email protected] 3Brigham Young University, Department of Biology and M. L. Bean Life Science Museum, Provo, Utah 84602 ­ [email protected]

Introduction species boundaries and molecular Our understanding of species diversity phylogenetic reconstructions of the in lichen­forming fungi is currently in a mycobiont suggests that one of the state of flux (Lumbsch and Leavitt, 2011). greatest challenges to accurate species There is mounting evidence indicating that identification of lichenized fungi is finding in many cases traditional taxonomy fails to and using appropriate character sets and accurately represent natural groups (ex.: analytical tools (Crespo and Pérez­Ortega, Caloplaca [Vondrák et al., 2009]; Letharia 2009; Printzen, 2009). [Kroken and Taylor, 2001]; and Few field lichenologists will be Rhizoplaca [Leavitt et al., 2011a]. While surprised to learn that a lichenized­fungus morphological and chemical character is difficult to identify. Xanthoparmelia differences have traditionally served as (Vainio) Hale is no exception and is proxies for identifying independent considered among the most notoriously evolutionary lineages (i.e. ‘species’), difficult taxonomic groups of character evolution in lichens is still macrolichens (Leavitt et al., 2011b). poorly understood, resulting in potentially Xanthoparmelia is the largest genus within confounding morphological and chemical the Parmeliaceae, with more than 800 taxonomic characters. It is not surprising recognized species (Amo de Paz et al., that the phenotypic expressions of 2010; Blanco et al., 2004; Hale, 1990). symbiotic systems, like lichens, may result This genus displays considerable in observable differences that do not morphological and chemical diversity; and necessarily reflect natural groups for slight morphological and/or chemical individual lichen symbionts (i.e. differences have often been used to mycobiont, photobiont, or any other distinguish species. Additionally, ‘biont’ associated with a lichen thallus). contrasting reproductive modes have also This incongruence between traditional been considered to provide important

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characters for diagnosing species within traditional Xanthoparmelia species Xanthoparmelia (Hale, 1990), although (Hodkinson and Lendemer, 2011; Leavitt many typically isidiate species also et al., 2011b; Thell et al., 2009). The clear produce apothecia on occasion. However, conflict between traditional species the use of molecular sequence data boundaries and conclusions drawn from suggests that chemical and morphological genetic data has resulted in a difficult characters traditionally used to define conundrum on how we should treat taxonomic groups within this genus have diversity within this important genus. been overemphasized (Amo de Paz et al., A sizeable portion of Xanthoparmelia 2010; Leavitt et al., 2011b; Leavitt et al., diversity is found in North America (Fig. 2011c). Adding to the taxonomic 1). Currently, a total of 85 species are confusion, morphologically cryptic recorded for North America, 35 of which species are commonly hidden within are known to occur in California. Many

Figure 1. Variation in morphology and habit within sampled Xanthoparmelia in western North America. (A) saxicolous attached taxon X. cumberlandia sensu lato (s. l.) with apothecia; (B) saxicolous attached taxon X. mexicana with isidia; (C) terricolous taxon X. wyomingica s. l., an intermediate growth­form between attached and vagrant forms; (D) vagrant taxon X. chlorochroa s. l.; (E) morphology of rare vagrant or semi­attached taxon X. idahoensis s. l.; (F) white­maculate upper cortex on X. camtschadalis s. l.; (G) lobe morphology and weakly maculate surface on X. stenophylla; (H) erhizinate lower surface of the vagrant taxon X. norchlorochroa s. l.; (I) rhizine characters on the vagrant taxon X. chlorochroa s. l. Photos by Steven Leavitt.

59 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Leavitt et al. ­ Xanthoparmelia species may be locally common to frequently reviewed (Crespo and abundant, but identification of even the Lumbsch, 2010; Crespo and Pérez­Ortega, most common species is often difficult. 2009; Lumbsch and Leavitt, 2011), it Morphologically and chemically diverse remains a controversial topic in species within Xanthoparmelia in western lichenology. Consequently, even the North America provide a useful model for interpretation and application of the term exploring the challenges associated with ‘cryptic species’ or ‘cryptic diversity’ integrating molecular sequence data with varies widely among users. We suggest traditional taxonomic characters to better that the most practical definition of understand species diversity in lichenized ‘cryptic’ species is simply where two or fungi. As part of our ongoing research, we more distinct species are erroneously have amassed over 4000 Xanthoparmelia classified (and hidden) under one species specimens from western North America name (Bickford et al., 2007). This does not and generated genetic sequence data for preclude the possibility that diagnostic more than 500 individuals. Here we briefly phenotypic characters supporting distinct report on our current findings and their species­level lineages may be identified in implications for understanding lichen future studies. Simply, the distinct species diversity in western North America. Using are unrecognizable using current results from our research on taxonomic characters. In some cases of Xanthoparmelia we specifically address cryptic diversity in lichenized fungi, a re­ two major issues currently facing examination of morphology and/or lichenologists: chemistry against a molecular (1) Cryptic diversity in ‘traditional’ phylogenetic hypothesis has revealed species and previously overlooked morphological (2) Polymorphic species representing and/or chemical characters supporting the multiple ‘traditional’ lichen fungal species separation of distinct phylogenetics clades within a single species­level lineage. as separate species (ex. Argüello et al., Here we synthesize the impact of these 2007; Divakar et al., 2010; Leavitt et al. two common phenomena in 2012; Spribille et al., 2011; Wirtz et al., Xanthoparmelia in western North America 2008). However, it has long been known and their implications for understanding by evolutionary biologists that distinct and identifying lichen species diversity. species do not need to have diagnosable morphological differences, (e.g. as Cryptic Species summarized by Mayr 1942; Mayr 1963). Dealing with multiple species­level Not surprisingly, within lichenized fungi lineages hidden under one species name is there are also cases in which no hardly limited to lichen symbionts. In fact, corroborating phenotypic characters have cryptic species have been commonly been identified supporting cryptic lineages documented across all forms of life, (Baloch and Grube, 2009; Leavitt et al., including bacteria, protists, plants, 2011a; Molina et al., 2011). animals, and other fungi. Although cryptic In our study of the genus diversity in lichen­forming fungi has been Xanthoparmelia in western North America

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we have found strong evidence of cryptic Similarly, specimens identifiable as the species (often recovered within isidiate species X. mexicana and X. plittii polymorphic lineages – discussed below) were each recovered in at least three within nearly all of the traditional species genetically distinct populations (Table 1). we have sampled (Fig. 2; Leavitt et al., Among the most surprising results of our 2011b; Leavitt et al., 2011c). For example, research, we found that the vagrant species the stictic acid containing species, X. X. chlorochroa was represented in least six cumberlandia, was found in at least four distinct species­level lineages (Table 1). distinct species­level lineages (Table 1). Overall, the pattern of cryptic diversity

Figure 2. A multilocus species tree, representing relationships between species­level genetic clusters inferred from four ribosomal loci and two protein coding markers generated from Xanthoparmelia specimens collected in western North America. Traditional combinations of morphological and chemical characters used for species recognition in Xanthoparmelia were not supported. Major diagnostic characters are distributed across divergent lineages, including (a) presence of isidia; (b) maculate upper cortex (in vagrant forms); (c) emaculate vagrant growth form; and (d) emaculate saxicolous forms. Underlined terminal labels indicate chemically polymorphic genetic clusters. Thickened branches are proportional to the level of posterior probability support (see Leavitt et al 2011b for complete details). Photos by Steven Leavitt.

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Table 1. Distribution of a total of 414 traditionally circumscribed species from western North America in species­level genetic clusters. Eight major genetic groups (A­H) were inferred from six genetic markers (ITS, IGS, group I intron, β­tubulin, and MCM7); and a total of 21 species­level genetic clusters (A1 – H2) were identified within these groups. Most inferred population clusters include multiple traditionally circumscribed species, and at the same time the most common morphological/chemical species are distributed in different population clusters. Values within each group represent the number of individuals recovered within each genetic cluster (see Leavitt et al. 2011b for complete details).

within traditional Xanthoparmelia species 1993; Rosentreter and McCune, 1992; was observed in 13 of the 19 currently Weber, 1977). Our findings clearly show accepted species sampled in connection that the most common traditional vagrant with our research (Leavitt et al., 2011b). species, X. camtschadalis and X. The results of our research suggest that chlorochroa, are actually composed of extensive morphological and chemical multiple distinct species­lineages within homoplasy (similarity arising from each group (Table 1), necessitating a evolutionary convergence) in careful rethinking of current taxonomic Xanthoparmelia tends to obscure concepts in vagrant forms of recognition of natural lineages within the Xanthoparmelia. These results provide mycobiont ­ whether based on thallus strong evidence for multiple independent morphology, medullary chemistry, or origins of vagrant forms in reproductive mode (Fig. 2). Evolutionary Xanthoparmelia in western North relationships between attached America; and both vagrant and attached (saxicolous) and vagrant growth­forms in forms may occur within a single species­ lichenized ascomycetes have long been level lineage. debated (Klement, 1950; Rosentreter, Uncertainty about the taxonomic

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significance of thallus morphology in Wirtz et al., 2008), while in other cases Xanthoparmelia has traditionally resulteAd specialized instruments, such as a in heavy reliance on medullary chemistry. scanning electron microscope (SEM), may The most common diagnostic secondary be required to observe diagnostic metabolites for the most Xanthoparmelia characters (see Pino­Bodas et al., 2012a), species in western North America are rendering the diagnostic character closely related β­orcinol depsidones (ex. practically unusable for most researchers. norstictic, salazinic, and stictic acids). This leads to an important point of Genetic, ecological, and biological factors discussion. Although corroborating influencing the expression of this highly morphological or chemical characters may characteristic group of compounds are support distinct lineages hidden under a unclear (Asplund and Gauslaa, 2007; traditionally circumscribed species, these Asplund et al., 2009; Solhaug et al., 2009). separate ‘species’ may remain practically Our research suggests that due to ‘cryptic’. Most lichenologists Dare independent changes in chemical charactCer experienced with taxonomy requiring a states, medullary chemistry has only hand lens, dissecting microscope, and limited value in delimiting natural groups chemical spot tests for accurate in Xanthoparmelia in western North identifications. In addition, the majority of America, at least as currently interpreted lichenologist likely have access to (Fig. 2). somewhat more sophisticated tools, such At this point in our research, we have as a compound microscope or thin­layer generally been unable to discern any chromatography; and accurate species morphological differences or striking identification using these more advanced geographic and/or ecological patterns approaches for identifying diagnostic among divergent groups with similar characters are standard for many lichen morphologies. However, additional a groups. However, the use of more posteriori examination of species­levEel specialized tools, such as SEM, Fhigh­ lineages inferred from genetic data may performance liquid chromatography yet reveal previously overlooked (HPLC), and polymerase chain reaction morphological, chemical and/or ecological (PCR) amplification of genetic data, is not characters supporting these lineages. A standard in the vast majority of taxonomic careful reexamination of putative ‘cryptic’ treatments. In spite of the fact that SEM, Xanthoparmelia lineages for corroborating HPLC, and genetic data can provide phenotypic characters (i.e. ecology, valuable characters for accurate species geography, morphology, and chemistry) identification, incorporating this data into will be important for developing an practical taxonomic treatments remains a integrative taxonomy and robust species challenge. boundaries supported by multiple lines of evidence. In some cases the corroborating Polymorphic Species phenotypic characters supporting cryptic Understanding the differences between lineages may be practically useful for field morphological variation within a species identification (ex. Divakar et al., 2010; and among closely related groups is

63 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Leavitt et al. ­ Xanthoparmelia central to identifying the diagnostic polymorphic species within this genus (ex. characters required for establishing the Cladonia gracilis group [Pino­Bodas accurate taxonomic boundaries. However, et al., 2011]; Cladonia arbuscula sensu in practice a clear demarcation between lato [Piercey­Normore et al., 2010]). intraspecific and interspecific variation is Species identification in the subject to observational bias and morphologically, biochemically, and individual interpretation. Accurate reproductively diverse lichen genus taxonomic circumscriptions may be Xanthoparmelia is notoriously confounded by varying levels of challenging. Our research on intraspecific variation among different Xanthoparmelia using molecular sequence species groups. While some species may data provides some insights into why have little variation, high levels of species identifications may be so difficult. intraspecific phenotyptic variation are In many cases, species­level genetic well­documented in some lichen­forming groups were shown to be morphologically fungi. A striking example is found in the and chemically polymorphic, containing terricolous cosmopolitan lichen­forming up to eight traditionally circumscribed species Cetraria aculeata, where in steppe Xanthoparmelia species within a single areas of Central Spain, Iran, and Ukraine species­level lineage (Table 1). For vagrant specimens display an exceptional example, one genetic cluster, ‘X. morphology that deviates from their isidiate/mix ’ D1 (Fig. 2) was dominated typical fruticose pattern (Pérez­Ortega et by four currently accepted species (X. al., 2012). Culberson et al. (1988) artfully dierythra [with norstictic acid]; X. demonstrated that in Cladonia neither mexicana [with salazinic acid]; X. plittii morphological tendencies nor distinct [with stictic acid]; and X. subplittii [with patterns of secondary metabolite stictic acid]), but also contained a vagrant production are necessarily correlated to specimen identified as X. chlorochroa and reproductive barriers among different non­isidiate specimens producing forms. In some cases, intraspecific abundant apothecia (Leavitt et al., 2011c). phenotypic variation within this genus has A similar pattern of high intraspecific been interpreted as evidence supporting morphological polymorphism is found in distinct species (Ahti, 2000), which may non­isidiate saxicolous and vagrant not reflect natural groups. For example, it Xanthoparmelia specimens common in the has recently been shown that several intermountain region of western North traditionally circumscribed species within America (Leavitt et al., 2011b). Again, the Cladonia gracilis group (C. using genetic data we identified distinct coniocraea, C. cornuta subsp. genetic clusters corresponding to groenlandica, and C. ochrochlora) likely ‘species’­level lineages each containing belong to a single morphologically the morphologically­circumscribed species polymorphic species (Pino­Bodas et al., X. chlorochroa (vagrant species containing 2011). In other cases in Cladonia, salazinic acid), X. coloradoënsis molecular sequence data supports (saxicolous species containing salazinic chemically and morphologically acid), and X. cumberlandia (saxicolous

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species containing stictic acid) (Fig. 2). chemical polymorphism may also occur Clearly, our results indicated that within species­level lineages in characters traditionally used to separate Xanthoparmelia. While these results Xanthoparmelia species are commonly highlight the limitations of using polymorphic within the population traditional taxonomic characters for clusters inferred from genetic data. separating natural groups within However, in most cases each distinct Xanthoparmelia, they also provide a population cluster showed a general trend valuable perspective on the importance of in the expression of secondary ongoing research in even the best­studied metabolites, suggesting at least some level lichen groups. of reproductive isolation between salazinic and stictic acid chemotypes. Where do we go from here? Intraspecific polymorphisms are even Accurate species circumscriptions and common in vagrant forms of identifications are essential for various Xanthoparmelia. Our results indicate that sub­disciplines of biology. The availability in one case a total of six traditionally of genetic data has allowed researchers to circumscribed vagrant species (X. identify species and to rigorously test chlorochroa, X. lipochlorochroa , X. species boundaries in lichen­forming fungi norchlorochroa , X. neochlorochroa, X. with a level of precision that was vagans, and X. wyomingica) were unimaginable a decade ago. In spite of our recovered within a single species­level progress with accurately recognizing genetic cluster, in addition to the diversity using molecular sequence data, saxicolous species X. coloradoënsis and X. successfully implementing these results cumberlandia (Table 1). Our research into a working taxonomy remains results indicate that a clear demarcation challenging. between intraspecific and interspecific While our results may contradict morphological and/or chemical variation traditional species boundaries within does not exist for a large proportion of Xanthoparmelia in western North Xanthoparmelia species in western North America, we are optimistic that our America (Fig. 2), in spite of general trends research represents substantial progress of morphological similarity within some towards a more accurate perspective on species­level lineages. species diversity in this important genus. Our research clearly indicates that our As a result of our research, the taxonomic current interpretation of morphological value of phenotypic characters in and chemical characters in Xanthoparmelia in western North America Xanthoparmelia is inadequate to is now better understood. For example, in accurately characterize species diversity. arid continental regions of western North As in other groups of lichen­forming America the vast majority of isidiate fungi, such as Bryoria (Velmala et al., Xanthoparmelia specimens belong to a 2009), Cladonia (Pino­Bodas et al., single species­level lineage, regardless of 2012b), Cetraria (Pérez­Ortega et al., whether the specimens produces salazinic, 2012), a high degree of morphological and stictic, or norstictic acid as the major

65 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Leavitt et al. ­ Xanthoparmelia extrolite. In spite of the morphological and within lichen groups where diagnostic chemical similarities of isidiate morphological characters are Xanthoparmelia specimens from more unidentifiable or practically not feasible. humid regions they all appear to belong to These are exciting times for completely distinct species­level lineages. lichenologists. A closer look at lichen Hence, understanding ecological and taxonomy, with the inclusion of new data, geographic patterns associated with the will help us to better understand the distinct genetic groups will be important in diversity of lichenized fungi, accurately developing a working taxonomy with interpret distribution patterns, and play a some Xanthoparmelia species. A second more important role in meaningful example includes Xanthoparmelia conservation practices. However, progress specimens collected from the will be accompanied by some level of Intermountain Area of western North uncertainty. In many taxonomic groups, America. In this region specimens including Xanthoparmelia, our traditional producing salazinic acid generally belong approach for species identification will to a lineage distinct from co­occurring likely need to be substantially modified. specimens producing stictic acid, in spite The search for corroborating of the high degree of variation in thallus morphological support for cryptic species morphology. In this case, species identified using molecular data will traditionally identified as ‘X. chlorochroa’, require meticulous and creative ‘X. coloradoënsis’, and ‘X. wyomingica’ approaches to assess phenotypic variation should likely be treated as a single in potentially unorthodox ways. We are morphological polymorphic species (all hopeful that lichenologists, who containing salazinic acid), separate from traditionally have been eager to include ‘X. cumberlandia’ (which produces stictic new methods, such as chromatography, in acid). However, it is important to keep in their routine identifications, will be mind that this is a region­specific pattern, amenable to include molecular techniques and each of these traditional species can be to their routine examination of specimens found in other distinct species­level for identification and classification. lineages. Although this may prove difficult to While we are strong advocates for the achieve by single individuals, especially application of independent data types in citizen scientists that traditionally play an developing an integrative taxonomy, there important role in lichen taxonomy (Poelt, is an increasing need to formally recognize 1992), the increasing number of the existence of phenotypically cryptic collaborative projects in the discipline species­level lineages in lichen­forming (e.g., Crespo et al., 2010; Gueidan et al., fungi (see Crespo and Lumbsch, 2010; 2009; Lumbsch et al., 2011) make us Crespo and Pérez­Ortega, 2009; Hibbett et optimistic that broad­scale collaborative al., 2011; Lumbsch and Leavitt, 2011). In approaches will facilitate the inclusion of some cases, a molecular taxonomy may molecular data in lichen research at all provide the most practical approach to levels. Integrating new data (including consistent treatment of mycobiont species novel morphological characters and

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genetic data) will be essential to the foliose lichen Lobaria pulmonaria. The accurately represent lichen diversity in Lichenologist 39: 273­278. Xanthoparmelia and other lichen groups. Asplund, J., Solhaug, K.A., & Y. Gauslaa. 2009. Hopefully, an improved perspective on Fungal depsidones ­ an inducible or constitutive lichen diversity also increases our defence against herbivores in the lichen Lobaria appreciation of these incredible symbiotic pulmonaria? Basic and Applied Ecology 10: 273­ systems. 278. Baloch, E. & M. Grube. 2009. Pronounced Acknowledgements genetic diversity in tropical epiphyllous lichen We are indebted to various colleagues fungi. Molecular Ecology 18: 2185­2197. for providing valuable material and field Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, assistance, notably J. Belnap, C. Björk, S. P.K.L., Meier, R., Winker, K., Ingram, K.K., Crawford, A. DeBolt, M. DeVito, R. Egan, and I. Das. 2007. Cryptic species as a window on T. Esslinger, M. Felix, R. Fuller, T. diversity and conservation. Trends in Ecology & Goward, T. Hardle, S. Hardle, J. Hertz, J. Evolution 22: 148­155. Hollinger, K. Knight, G. Lind, the Leavitt Blanco, O., Crespo, A., Elix, J.A., Hawksworth, family, J. Marsh, J. Muscha, B. McCune, D.L., & H.T. Lumbsch. 2004. A Molecular M. Robinson, R. Rosentreter, G. Shrestha, Phylogeny and a New Classification of Parmelioid and T. Wheeler. We also thank T. Goward Lichens Containing Xanthoparmelia­Type and T. Lumbsch for invaluable discussion. Lichenan (Ascomycota: Lecanorales). Taxon 53: We gratefully acknowledge support from 959­975. the California Lichen Society Education Crespo, A., Kauff, F., Divakar, P.K., del Prado, Grants Program, The Ruth L. Glacy R., Perez­Ortega, S., Amo de Paz, G., Foundation, and the Brigham Young Ferencova, Z., Blanco, O., Roca­Valiente, B., University Office of Research and Nunez­Zapata, J., Cubas, P., Arguello, A., Elix, Creative Activities. J.A., Esslinger, T.L., Hawksworth, D.L., Millanes, A., Molina, M.C., Wedin, M., Ahti, T., Literature Cited & A. Aptroot, et al. 2010. Phylogenetic generic Ahti, T., 2000. Cladoniaceae Flora Neotropical classification of parmelioid lichens (Parmeliaceae, Monograph 78: 1­362. Ascomycota) based on molecular, morphological Amo de Paz, G., Lumbsch, H.T., Cubas, P., Elix, and chemical evidence. Taxon 59: 1735­1753. J.A., & A. Crespo. 2010. The genus Karoowia Crespo, A. & H. T. Lumbsch. 2010. Cryptic (Parmeliaceae, Ascomycota) includes unrelated species in lichen­forming fungi. IMA Fungus 1: clades nested within Xanthoparmelia. Australian 167­170. Systematic Botany 23: 173­184. Crespo, A. & S. Pérez­Ortega. 2009. Cryptic Argüello, A., Del Prado, R., Cubas, P., & A. species and species pairs in lichens: A discussion Crespo. 2007. Parmelina quercina (Parmeliaceae, on the relationship between molecular phylogenies Lecanorales) includes four phylogenetically and morphological characters. Anales del Jardin supported morphospecies. Biological Journal of Botanico de Madrid 66(S1): 71­81. the Linnean Society 91: 455­467. Culberson, C.F., Culberson, W.L., & A. Asplund, J. & Y. Gauslaa. 2007. Content of Johnson. 1988. Gene flow in lichens. American secondary compounds depends on thallus size in Journal of Botany 75: 1135­1139.

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Divakar, P.K., Figueras, G., Hladun, N., & A. Porter, L.D., Johnson, L.A., & L.L. St. Clair. Crespo. 2010. Molecular phylogenetic studies 2011a. Complex patterns of speciation in reveal an undescribed species within the North cosmopolitan ‘‘rock posy’’ lichens – Discovering American concept of Melanelixia glabra and delimiting cryptic fungal species in the lichen­ (Parmeliaceae). Fungal Diversity 42: 47­55. forming Rhizoplaca melanophthalma species­ Gueidan, C., Savi, S., Thues, H., Roux, C., complex (Lecanoraceae, Ascomycota). Molecular Keller, C., Tibell, L., Prieto, M., Heimarsson, S., Phylogenetics and Evolution 59: 587­602. Breuss, O., Orange, A., Froberg, L., Wynns, Leavitt, S.D., Johnson, L.A., Goward, T., & L.L. A.A., Navarro­Rosines, P., Krzewicka, B., St. Clair. 2011b. Species delimitation in Pykaelae, J., Grube, M., & F. Lutzoni. 2009. taxonomically difficult lichen­forming fungi: An Generic classification of the Verrucariaceae example from morphologically and chemically (Ascomycota) based on molecular and diverse Xanthoparmelia (Parmeliaceae) in North morphological evidence: recent progress and America. Molecular Phylogenetics and Evolution remaining challenges. Taxon 58: 184­208. 60: 317­332. Hale, M.E. 1990. A synopsis of the lichen genus Leavitt, S.D., Johnson, L., & L.L. St. Clair. Xanthoparmelia (Vainio) Hale (Ascomycotina, 2011c. Species delimitation and evolution in Parmeliaceae). Smithsonian Institution Press, morphologically and chemically diverse Washington D.C. communities of the lichen­forming genus Hibbett, D.S., Ohman, A., Glotzer, D., Nuhn, M., Xanthoparmelia (Parmeliaceae, Ascomycota) in Kirk, P., & R. H. Nilsson. 2011. Progress in western North America. American Journal of molecular and morphological taxon discovery in Botany 98: 175­188. Fungi and options for formal classification on Lumbsch, H.T., Ahti, T., Altermann, S., Amo de environmental sequences. Fungal Biology Reviews Paz, G., Aptroot, A., Arup, U., Barcenas Peña, 25: 38­47. A., Bawingan, P.A., Benatti, M.N., Betancourt, Hodkinson, B.P. & J. C. Lendemer. 2011. L., Björk, C.R., Boonpragob, K., Brand, M., Molecular analyses reveal semi­cryptic species in Bungartz, F., Caceres, M.E.S., Candan, M., Xanthoparmelia tasmanica. Bibliotheca Chaves, J.L., Clerc, P., Common, R., Coppins, Lichenologica 106, 108­119. B.J., Crespo, A., Dal Forno, M., Divakar, P.K., Klement, O. 1950. Über die Artberechtigung Duya, M.V., Elix, J.A., Elvebakk, A., einiger Parmelien. Berichten der Deutschen Fankhauser, J., Farkas, E., Ferraro, L.I., Botanischen Gesellschaft, Jahrgang 63: 47­52. Fischer, E., D.J., G., Gaya, E., Giralt, M., Kroken, S. & J. W. Taylor. 2001. A Gene Goward, T., Grube, M., Hafellner, J., Genealogical Approach to Recognize Phylogenetic Hernandez, J.E., Herrera­Campos, M.A., Kalb, Species Boundaries in the Lichenized Fungus K., Kärnefelt, I., Kantvilas, G., Killmann, D., Letharia. Mycologia 93: 38­53. Kirika, P., Knudesn, K., Komposch, H., Leavitt, S.D., Esslinger, T.L., & H.T. Lumbsch. Kondratyuk, S., Lawrey, J.D., Mangold, A., 2012. Neogene­dominated diversification in Marcelli, M.P., McCune, B.P., Michlig, A., neotropical montane lichens: dating divergence Miranda Gonzalez, R., Moncada, B., Naikatini, events in the lichen­forming fungal genus A., Nelsen, M.P., Øvstedal, D.O., Palice, Z., Oropogon (Parmeliaceae). American Journal of Papong, K., Parnmen, S., Pérez­Ortega, S., Botany doi:10.3732/ajb.1200146. Printzen, C., Rico, V.J., Rivas Plata, E., Robayo, Leavitt, S.D., Fankhauser, J.D., Leavitt, D.H., J., Rosabal, D., Ruprecht, U., Salazar Allen, N.,

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Sancho, L., Santos de Jesus, L., Santos Vieira, Cladonia gracilis group (Cladoniaceae, T., Schultz, M., Seaward, M.R.D., Sérusiaux, E., Ascomycota). Organisms Diversity & Evolution Schmitt, I., Sipman, H.J.M., Sohrabi, M., 11: 343­355. Søchting, U., Søgaard, M.Z., Sparrius, L.B., Pino­Bodas, R., Burgaz, A.R., Martin, M.P., & Spielmann, A., Spribille, T., Sutjaritturakan, J., H.T. Lumbsch. 2012a. Species delimitations in the Thammathaworn, A., Thell, A., Thor, G., Thüs, Cladonia cariosa group (Cladoniaceae, H., Timdal, E., Truong, C., Türk, R., Umaña Ascomycota). The Lichenologist 44: 121­135. Tenorio, L., Upreti, D., van den Boom, P., Vivas Pino­Bodas, R., Martín, M., & A. Burgaz. Rebuelta, M., Wedin, M., Will­Wolf, S., Wirth, 2012b. Cladonia subturgida and C. iberica V., Wirtz, N., Yahr, R., Yeshitela, K., Ziemmeck, (Cladoniaceae) form a single, morphologically and F., Wheeler, T., & R. Lücking. 2011. One hundred chemically polymorphic species. Mycological new species of lichenized fungi: a signature of Progress 11: 269­278. undiscovered global diversity. Phytotaxa 18: 1­ Poelt, J. 1992. Lichenologie und Lichenologen in 127. in Bayern eine Reverenz vor den Amateuren. Lumbsch, H.T. & S. D. Leavitt. 2011. Goodbye Hoppea 50: 527­536. morphology? A paradigm shift in the delimitation Printzen, C. 2009. Lichen Systematics: The Role of species in lichenized fungi. Fungal Diversity 50: of Morphological and Molecular Data to 59­72. Reconstruct Phylogenetic Relationships. Progress Mayr, E. 1942. Systematics and the origin of in Botany 71. Springer Berlin Heidelberg, pp. 233­ species from the viewpoint of a zoologist. Harvard 275. University Press, Cambridge. Rosentreter, R. 1993. Vagrant Lichens in North Mayr, E. 1963. Animal species and evolution. America. The Bryologist 96: 333­338. Harvard University Press, Cambridge. Rosentreter, R. & B. McCune, B. 1992. Vagrant Molina, M., Del­Prado, R., Divakar, P., Dermatocarpon in Western North America. The Sánchez­Mata, D., & A. Crespo. 2011. Another Bryologist 95: 15­19. example of cryptic diversity in lichen­forming Solhaug, K.A., Lind, M., Nybakken, L., & Y. fungi: the new species Parmelia mayi Gauslaa. 2009. Possible functional roles of (Ascomycota: Parmeliaceae). Organisms Diversity cortical depsides and medullary depsidones in the & Evolution 11: 331­342. foliose lichen Hypogymnia physodes. Flora ­ Pérez­Ortega, S., Fernández­Mendoza, F., Morphology, Distribution, Functional Ecology of Raggio, J., Vivas, M., Ascaso, C., Sancho, L.G., Plants 204: 40­48. Printzen, C., & A. de los Ríos. 2012. Extreme Spribille, T., Klug, B., & H. Mayrhofer. 2011. A phenotypic variation in Cetraria aculeata phylogenetic analysis of the boreal lichen (lichenized Ascomycota): adaptation or incidental Mycoblastus sanguinarius (Mycoblastaceae, modification? Annals of Botany 109: 1133­1148. lichenized Ascomycota) reveals cryptic clades Piercey­Normore, M.D., Ahti, T., & T. Goward. correlated with fatty acid profiles. Molecular 2010. Phylogenetic and haplotype analyses of four Phylogenetics and Evolution 59: 603­614. segregates within Cladonia arbuscula s.l. Botany Thell, A., Elix, J.A., & U. Søchting. 2009. 88: 397­408. Xanthoparmelia lineola s. l. in Australia and North Pino­Bodas, R., Burgaz, A., Martín, M., & H. T. America. Bibliothecia Lichenologica 99: 393­404. Lumbsch. 2011. Phenotypical plasticity and Velmala, S., Myllys, L., Halonen, P., Goward, T., homoplasy complicate species delimitation in the & T. Ahti. 2009. Molecular data show that Bryoria

69 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Leavitt et al. ­ Xanthoparmelia fremontii and B. tortuosa ( Parmeliaceae) are Wirtz, N., Printzen, C., & H. T. Lumbsch. 2008. conspecific. The Lichenologist 41: 231­242. The delimitation of Antarctic and bipolar species of Vondrák, J., Říha, P., Arup, U., & U. Søchting. neuropogonoid Usnea (Ascomycota, Lecanorales): 2009. The taxonomy of the Caloplaca citrina a cohesion approach of species recognition for the group ( Teloschistaceae) in the Black Sea region; Usnea perpusilla complex. Mycological Research with contributions to the cryptic species concept in 112: 472­484. lichenology. The Lichenologist 41: 571­604. Weber, W.A., 1977. Environmental Modifications. In: Seaward, M.R.D. (Ed.), Lichen Ecology. Academic Press, New York, pp. 9­29.

Xanthoparmelia aff. cumberlandia, Aquarius Plateau, Utah, USA. Photo by Steven Leavitt.

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Survey of the Lichen Flora of Palos Verdes, Southern California

Jason Hollinger 11034 White Oak Road, Waynesville, NC 28785 [email protected]

A preliminary lichen survey of the work on saxicolous communities is Palos Verdes Peninsula has yielded 64 needed. Bluff Cove is identified as a species, 57 of which are here reported as promising site for further study. new to the area. Of these, 16 are shown to be substantially more abundant on the Introduction Channel Islands than on the mainland, Palos Verdes Peninsula juts into the while 5 appear to be more common on the ocean in southern Los Angeles County mainland. Six ­ Caloplaca stipitata between San Pedro and Redondo Beach Wetmore, Dendrographa leucophaea (Figure 1). It is justly famous for its (Tuck.) Darb., Lecidella scabra (Taylor) dramatic, nearly continuous ring of sea Hertel & Leuckert, Niebla ceruchis Rundel cliffs. Its wild, rugged prospect stands in & Bowler, Niebla combeoides (Nyl.) stark contrast to the nearby Los Angeles Rundel & Bowler and Roccella gracilis basin, one of the most heavily urbanized Bory ­ are rarely reported from mainland and fastest growing regions in North southern California; their occurrence America with a population of nearly 18 adjacent to one of the most densely million. Only in recent decades has populated areas in the U.S. is noteworthy. suburban development begun to encroach, The richest lichen communities occur on especially in the western portions. coastal sage and chaparral on north­ and Like many other mountain ranges in northwest­facing slopes, although further southern California, Palos Verdes Peninsula is an uplifted fault block. Roughly 1 to 2 million years ago it rose from the sea as an island, effectively part of the Channel Islands chain. Only toward the end of the Pleistocene, about 10 to 20 thousand years ago, did infilling of the Los Angeles basin finally connect it with the mainland (Byhower 2007). The rocks here consist primarily of 4 to 16 million year old sediments of the Monterey Figure 1. Map of southern California, showing Palos Formation (Altamira Shale, Mulaga Verdes (star) in relation to the Channel Islands, Santa Mudstone and Valmonte Diatomite) Barbara (SB), Santa Monica Mountains (SM), overlying the much older Catalina Schist. Orange County (OC), San Jacinto Mountains (SJ), The Monterey Formation in this area is San Diego (SD), and urban and agricultural lands geologically complex, encompassing large (dark areas). quantities of siliceous, carbonaceous and

71 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Hollinger ­ Palos Verdes phosphatic sediments and extensive tuff subsequently spreading to the Santa beds in the sea cliffs and hills above, as Monica Mountains and Orange County well as basalt intrusions often exposed at (Wells & Baptista 1979). their base (Conrad & Ehlig 1987). Owing to the inclination of the strata and the Previous Work chemical nature of the underlying Notwithstanding its proximity to Los sediments, portions of the southern half of Angeles, Palos Verdes has received little the peninsula are effectively enormous, attention from lichenologists. There are no slow­moving landslides (Morris 2000). It lichenological papers dealing specifically is this that has prevented widespread with the area. A search of CNALH housing development here. The slope at (http://lichenportal.org/portal) yielded only Portuguese Bend, for example, is presently 41 collections: 9 by H.E. Hasse between subsiding at a rate of approximately 1 to 2 1895 and 1912, one by W.A. Setchell in m per year (Calabro et al. 2010). 1896 and one by A.C. Herre in 1905, all Vegetationally, the coast of the peninsula from San Pedro, and 12 by W.A. Weber in is dominated by the Coastal Sage Scrub 1952 from the hills above Portuguese community (O’Leary 1990). Common Bend and Abalone Cove. Between 1908 shrubs include Artemisia californica, and 1914, Hasse made 17 additional Salvia leucophylla, S. mellifera, Lycium collections at nearby Clifton­by­the­Sea in californicum and Atriplex californica. southern Redondo Beach (Figure 2). Since Larger chaparral species like Rhus many of these are saxicolous and are listed integrifolia and Quercus spp. occur locally on northwest­facing slopes. Several canyons with perennial streams support riparian communities, including shrubs like Salix lasiolepis and Sambucus mexicana. Occasional small fires have occurred throughout the area. It is now well established that the flora and fauna of Palos Verdes Peninsula have their closest affinities with the Channel Islands. Several plants and animals occur nowhere else on the mainland. Examples include the plants Dudleya virens and Crossosoma californicum, (Munz 1970, Baldwin et al. 2012) and the Dusky Figure 2. Map of Palos Verdes, showing locations of Orange­crowned Warbler (also known to historical records: (A) Clifton­by­the­Sea, (B) breed at Point Loma, San Diego; Bradley Abalone Cove, (C) Portuguese Bend and (D) White 1980). The sedentarius subspecies of Point. Also shown are the four localities visited in Allen’s Hummingbird, formerly endemic connection with the present study: (1) Bluff Cove, to the Channel Islands, began to colonize (2) Paseo Lunado, (3) Point Vincente and (4) Sacred the mainland at Palos Verdes in the 1960s, Cove.

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as simply “near Clifton” it seems likely that they were actually collected in Palos Verdes, the only rocky cliffs near Redondo Beach. Consequently, I have included these records in the present study. In contrast to the meager work at Palos Verdes, other lichen hotspots in southern California (Figure 1) have received Figure 3. Bluff Cove, looking south. Photo by Jason considerable attention from lichenologists. Hollinger. To be sure, no fewer than 13 checklists have appeared in the last two decades: the of which 23 were epiphytes and the Santa Monica Mountains (Knudsen 2005, remaining 11 saxicoles. I attribute this rich 2007, Knudsen et al. 2008, Knudsen & flora to the gentle, north­facing bluffs Kocourková 2009b, 2010b), the coast of which support a uniquely well developed Orange and San Diego Counties (Bratt sage scrub community. 1997, Knudsen et al. 2008, Knudsen & As elsewhere along the coast of Palos Kocourková 2009a), the San Jacinto Verdes, the ocean spray zone is essentially Mountains (Knudsen & Kramer 2007), devoid of lichens, presumably reflecting Santa Barbara (Tucker 2010), and the the highly erodible nature of the rocks, Channel Islands (Bowler et al. 1996, Bratt here mostly shale and mudstone. Much 1993, 1999, Knudsen 2008, 2009, more productive are the stable basalt sills Knudsen & Kocourková 2010a, 2012). exposed just above the spray zone. Here I Taken together these papers report found extensive gray­brown patches of approximately 900 species of lichens, Lecania fructigena and orange colonies of lichenicolous fungi and allied fungi from Caloplaca luteominia var. luteominia, the the region. latter occasionally present also on stable soil. Fieldwork The crumbling, almost soil­like breccia On 18 January 2012, I paid a brief visit in shaded gullies supports a number of to Palos Verdes. Prior to my foray I interesting species, including Verrucaria selected four promising sites (numbered mimicrans, V. viridula, Buellia abstracta locations on Figure 2). Each of these sites and the sorediate Lecidella scabra. It is was carefully surveyed and representative worth noting that during my visit, these samples of all species encountered were well­sheltered microsites retained dew collected. Below I give a brief summary of well into the day. The southwest endemic each location with notes on the lichens species Cladonia nashii is locally encountered. abundant here. The richest lichen communities, Bluff Cove however, occur on the thick scrub, which By far the richest lichen flora was seen support an epiphytic flora characteristic of in Bluff Cove, on the northwestern corner coastal southern California, including of the peninsula, where I found 34 species, Cliostomum griffithii, Diploicia canescens,

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Flavoparmelia caperata, Niebla cephalota, Pyrrhospora quernea and Ramalina spp. Of particular interest is a population of Niebla ceruchis, now apparently rare in mainland southern California, where it has been collected recently only at Point Loma in San Diego (Bratt 4312 (SBBG), Knudsen 3257, 10009 (UCR)), Conejo Mountain in the Santa Monica Mountains (Knudsen 3950 (UCR)), and Santa Barbara (Nash 11203 (ASU)). This species is much more common in San Luis Obispo and Figure 4. Looking north from Point Vincente. Photo Monterey Counties (numerous specimens by Jason Hollinger. at SBBG) and Baja California. On the Channel Islands it is quite common southern California (but more common on (Knudsen & Kocourková 2012). the Channel Islands). This is an extraordinarily variable species (Figure 5). Arroyo at top of Paseo Lunado The apothecia vary from pale pink or This long gully is accessed from the top yellow to purple­brown or almost black of Paseo Lunado. The perennial (often multicolored), while the thallus watercourse is choked with willow, while ranges from thin and continuous to thick the surrounding slopes are open grassland and wrinkled, sometimes almost with scattered live oaks. The lichen flora dissolving into granular areoles or soredia here is extremely impoverished. Extensive (Ekman 2004). searching turned up only two saxicolous specimens of Caloplaca, although I noted Sacred Cove scattered unidentifiable fragments of This small cove near Portuguese Bend various terricolous and epiphytic lichens, on the southern coast of the peninsula has perhaps damaged by fire. been set aside as an ecological reserve. The cliffs here are continuously slumping, Cliff tops north of Point Vincente resulting in gentle south­facing slopes. Here as elsewhere along the top of the Sage scrub occurs sporadically throughout sea cliffs, the native vegetation is the area. A small pine grove covers a knoll dominated by an arid, windswept sage some distance back from the western community. These low shrubs support a headland, an unusual forest community for limited epiphytic crustose flora, but Palos Verdes. apparently little else. More might be found Of the eight species collected in this on the cliffs below. location, all crustose, five were saxicolous Of the three species I collected here, and three epiphytic. Sigridea californica is most notable is Bacidina californica, a particularly abundant on rough old bark of relatively rare lichen on the mainland of various shrubs. The unlichenized Arthonia

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Figure 5: Variation in apothecia and thalli of Bacidina californica. A: Palos Verdes, Hollinger 4424; B: San Gabriel Mountains, Hollinger 4528; C: isotype at UBC. Scale bar is 1 mm. Photos A & B by Jason Hollinger. Photo C by Curtis Björk.

albopulverea is locally common on both within brackets) are considered to be smooth­ and rough­barked stems. The erroneous or questionable. apothecia of the latter species are *Arthonia albopulverea Nyl. — Bluff remarkably variable (Figure 6). It forms a Cove: on limb of Rhus in chaparral, thin, white, unlichenized stain on living or Hollinger 4399, 4407; on stalk of dead dead bark with thin, black, epruinose herbaceous plant near beach, Hollinger apothecia. The apothecia vary from 4410; Sacred Cove: on limb of Pinus, subglobose to strongly asterisk­shaped. Hollinger 4420; on limb of dead unknown The spores are septate to finally exotic shrub, Hollinger 4428, 4431; all submuriform, but care must be taken to det. by K. Knudsen. find mature spores free of the ascus. Arthonia lecanactidea Zahlbr. — White Point: 1895, Hasse 2359 (MIN), this is the Annotated Checklist type specimen; on Lycium californicum, The material collected for this study, 1897 or 1899, Hasse 1316 (MICH, MIN), combined with historical records and isotypes. reports, brings the lichen flora for Palos Arthonia pruinata (Pers.) Steudel ex A.L. Verdes Peninsula to 64 species. Names Sm. — Reported as A. impolita (Ehrh.) marked with an asterisk (*) are new Borr. var. impolita and var. reports. Several historical records (shown chiodectonoides Tuck. by Hasse (1913) on

Figure 6. Variation in apothecia of Arthonia albopulverea at Palos Verdes. A: Hollinger 4407, B: Hollinger 4399, C­D: Hollinger 4431, E: Hollinger 4420. Scale bar is 1 mm. Photos by Jason Hollinger.

75 BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 19(2), 2012 Hollinger ­ Palos Verdes shrubs from White Point. — Hasse (1913) reports this as Blastenia *Bacidina californica S. Ekman — Point ferruginea (Huds.) Arnold var. fraudans T. Vincente: on twig of cfr. sage, Hollinger Fries on arenaceous shale at White Point. 4424; on dead twig of Cupressus Probably based on a misidentification macrocarpa, Hollinger 4426, det. by (Tucker 2007).] Knudsen. The apothecia were black with *Caloplaca luteominia (Tuck.) Zahlbr. dark brown exciple, dark brown to slightly var. luteominia — Clifton­by­the­Sea: greenish epihymenium, hyaline hymenium 1912, Hasse s.n. (ASU, MIN 865893, and dark brown hypothecium; the spores 865894), det. by C. Wetmore; Bluff Cove: were hyaline, fusiform, mostly 5­septate, abundant on outcrops above spray zone, ~12­13 x 2.5­3 μm, both ends equally Hollinger 4403a; on soil under lip at edge tapered; the pycnidia were in steep­sided of cliff, Hollinger 4402; Paseo Lunado: on verrucae with conidia ~2.5 x 1 μm. small rock on ground, Hollinger 4422b; *Buellia abstracta (Nyl.) H. Olivier — Sacred Cove: on rotten rock on ground in Bluff Cove: on crumbling rock in gully, pine grove, Hollinger 4436b. The thallus Hollinger 4414. The name B. sequax varied from entirely lacking to well­ (Nyl.) Zahlbr. was misapplied to this developed, beige and rimose. species in the Sonoran Flora (Knudsen, *Caloplaca pyracea (Ach.) Th. Fr. — pers. comm.). Bluff Cove: common on twigs of dead *Buellia alboatra (Hoffm.) Th. Fr. (= sage in chaparral, Hollinger 4396; on Diplotomma alboatrum (Hoffm.) Flotow) exposed dead rhizome of unknown plant, — Portuguese Bend: on rock, 24 Dec. Hollinger 4395b. In the past the name C. 1952, Weber S­986b (ASU). holocarpa (Hoffm. ex Ach.) A.E. Wade *Buellia halonia (Ach.) Tuck. — has been widely applied to epiphytic Portuguese Bend: on chert, 24 Dec. 1952, material of this taxon with orange Weber S­996 (ASU), det. by F. Bungartz in apothecia and scant orange thallus, but is 2003. now restricted to a saxicolous species not *Buellia maritima (A. Massal.) Bagl. — yet reported from California (Arup 2009). Portuguese Bend: on rock, 24 Dec. 1952, *Caloplaca stanfordensis H. Magn. — Weber S­986a (ASU), as B. subalbula Clifton­by­the­Sea: on twig, May 1914, (Nyl.) Müll. Arg., a misapplied name Hasse Exs. 207 (ASU), as C. cerina (Knudsen, pers. comm.). (Ehrh.) Zahlbr.; Bluff Cove: common on *Buellia punctata (Hoffm.) A. Massal. (= twigs of dead sage, Hollinger 4430. Less Amandinea punctata (Hoffm.) Coppins & common than the previous species; Scheid.) — Bluff Cove: on trunk of Rhus distinguished by the gray outer thalline in chaparral, Hollinger 4411; Point rims. Tucker (2007) reports a duplicate Vincente: on dead twig of Cupressus specimen at SBBG as C. cerina, however macrocarpa, Hollinger 4425. the one at ASU was annotated C. *Caloplaca bolacina (Tuck.) Herre — stanfordensis. Wetmore (2007) shows a Sacred Cove: on concrete wall (not dot for Palos Verdes on the map for the collected). latter, but not the former. [Caloplaca fraudans (Th. Fr.) H. Olivier *Caloplaca stipitata Wetmore — Bluff

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Cove: on twigs of dead sage in chaparral, *Diploicia canescens (Dickson) A. Hollinger 4397, juvenile specimen, det. by Massal. — Clifton­by­the­Sea: on dead K. Knudsen. Sambucus, 5 May 1908, Hasse s.n. (MIN *Caloplaca subsoluta (Nyl.) Zahlbr. — 875449, MSC 114146); Bluff Cove: Paseo Lunado: on small rock on ground, common on twigs of dead sage, Hollinger Hollinger 4422a. 4390. The Hasse specimen at MSC was *Chrysothrix xanthina (Vainio) Kalb — determined by H. Imshaug in 1982. See Bluff Cove: on sheltered twigs of dead also notes under Pyxine sorediata. sage in chaparral, Hollinger 4392. *Diploschistes scruposus (Schreber) *Cladonia nashii Ahti — Bluff Cove: Norman — Portuguese Bend: on chert and locally common on moss and soil under sandstone, 24 Dec. 1952, Weber 992 chaparral, Hollinger 4413, det. by K. (ASU). Knudsen. Very similar to C. hammeri Ahti *Evernia prunastri (L.) Ach. — Bluff which is also common in the chaparral of Cove: on twigs of dead sage (not southern California. Cladonia nashii is collected). distinguished by the presence of true *Flavoparmelia caperata (L.) Hale — soredia and atranorin (best revealed by a Bluff Cove: on twigs of dead sage (not drop of KOH in the cups, although TLC is collected). sometimes required; Knudsen, pers. *Lecania brunonis (Tuck.) Herre — comm.). White Point: Sep. 1912, Hasse Exs. 36 *Cliostomum griffithii (Sm.) Coppins — (ASU), det. by P.P.G. van den Boom in Bluff Cove: on bark and twigs of Rhus in 2003. chaparral, Hollinger 4400; Sacred Cove: Lecania dudleyi Herre — White Point: on limb of dead unknown tree, Hollinger Sep. 1912, Hasse Exs. 36b (ASU), det. by 4427. B.D. Ryan in 1999. Reported on *Cresponea chloroconia (Tuck.) Egea & calcareous rocks at White Point by Hasse Torrente — San Pedro: on dead twigs of (1913). Rhus, Nov. 1895, Hasse s.n. (ASU Lecania franciscana (Tuck.) K. Knudsen 683703). Probably originally recorded as a & Lendemer — Abalone Cove: 24 Dec. Lecanactis, but ASU records it as 1952, Weber 18093, L­1019, as Catillaria Cresponea premnea (Ach.) Egea & franciscana (Tuck.) Herre; Bluff Cove: on Torrente. Reports of C. premnea are outcrop along beach, Hollinger 4408. This apparently misidentifications of C. is a creamy­whitish, scurfy crust with chloroconia (Egea et al. 2004a). mostly 1­septate spores, 16­22 x 4.5­5 μm. Dendrographa leucophaea (Tuck.) Darb. Reported for White Point by Hasse (1913) — White Point: on twigs, Dec. 1896, as Catillaria franciscana. See Knudsen & Setchell 5125 (MIN); 1898, Hasse s.n. Lendemer 2007 for taxonomic notes. See (MIN 664458, 900249). Reported by notes under L. subdispersa (Nyl. ex B.D. Hasse (1913). Ryan) B.D. Ryan in van den Boom & *Dimelaena radiata (Tuck.) Müll. Arg. — Ryan 2004 for comments on Hasse’s Bluff Cove: on outcrop along beach, collections. Hollinger 4409. *Lecania fructigena Zahlbr. — Bluff

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Cove: abundant on outcrops along beach, H.T. Lumbsch in 2002. Hollinger 4401, 4403b. This is an *Lecanora hagenii (Ach.) Ach. — Bluff irregularly verrucose to subsquamulose, Cove: on exposed dead rhizome, Hollinger grayish to brownish crust with indistinctly 4395a. Very similar to L. dispersa, and septate spores, 10­15 x 4­5 μm and both can occur on either bark or rock, but abundant pycnidia with conidia 14­24 x 1 this species is distinguished by lack of μm. POL+ granules in epihymenium and *Lecania pacifica Zahlbr. ex B.D. Ryan & amphithecial cortex. van den Boom — Clifton­by­the­Sea: on *Lecidella scabra (Taylor) Hertel & rock, 1914, Hasse L­79350, as L. Leuckert — Bluff Cove: directly on rock angelensis Zahlbr. nom. prov. Can be very in chaparral gully, Hollinger 4417, det. by similar to L. brunonis and L. fructigena, K. Knudsen. This is a greenish­yellowish, see van den Boom & Ryan 2004. nearly leprose crust with soredia tending *Lecanographa subdryophila (Follmann to clump; C+ reddish orange, KC+ orange, & Vězda) Egea & Torrente — Clifton­by­ UV+ orange. the­Sea or San Pedro: 1895, Hasse s.n. *Leprocaulon microscopicum (Vill.) W. (ASU), det. by Egea & Torrente in 1994. Gams ex D. Hawksw. — Abalone Cove: Lecanora crenulata Hooker — Clifton­ on soil and rock just above tide mark, 24 by­the­Sea: 1914, Hasse s.n. (MIN Dec. 1952, Weber 1018 (ASU), det. by 901175); Portuguese Bend: on chert and B.D. Ryan in 1999. Knudsen & sandstone, 24 Dec. 1952, Weber S­997 Kocourková (2012) call this L. (ASU). Both originally recorded as L. americanum Lendemer & Hodkinson, hagenii, revised by Śliwa (2007). ined. *Lecanora dispersa (Pers.) Sommerf. — *Mobergia angelica (Stizenb.) H. Bluff Cove: on mossy outcrop in chaparral Mayrhofer & Sheard — Clifton­by­the­ gully, Hollinger 4416; Sacred Cove: on Sea: 1914, Hasse s.n. (ASU), det. by H. rotten rock on ground in grove, Hollinger Mayrhofer in 2002. 4436a. All spot tests negative for my *Naetrocymbe punctiformis (Schrank) material, more commonly with P+ R.C. Harris — Bluff Cove: on twigs of orange/red rims. See also L. hagenii. dead sage, Hollinger 4406; Point *Lecanora expallens Ach. — Bluff Cove: Vincente: on twig of cfr. sage, Hollinger on Rhus limb, Hollinger 4391, det. by J. 4423; both det. by K. Knudsen. This is an Lendemer. Superficially resembling a undercollected species. It is an Lepraria, with a finely powdery leprose unlichenized, thin, whitish crust with thallus, however very close inspection hemispherical, black, epruinose apothecia revealed a few minute fragments of intact and 1­septate, hyaline, slightly asymmetric gray­corticate thallus. Usnic and spores, 18­22 x 5­6 µm. thiophanic acids are present (C+ and KC+ *Niebla cephalota (Tuck.) Rundel & orange). Bowler — Bluff Cove: locally common on *Lecanora gangaleoides Nyl. — twigs of dead sage, Hollinger 4385. Portuguese Bend: on chert and sandstone, *Niebla ceruchis Rundel & Bowler — 24 Dec. 1952, Weber S­988 (ASU), det. by Bluff Cove: one fair population on twigs

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of dead sage, Hollinger 4384. [Pyxine sorediata (Ach.) Mont. — Clifton­ *Niebla combeoides (Nyl.) Rundel & by­the­Sea: on dead Sambucus, 1908, Bowler — San Pedro: on granite sea cliff, Hasse L­74581 (ASU). Misidentification 28 May 1905, Herre 510 (SBBG). Also of Diploicia canescens (Tucker 2007). reported tentatively by Hasse (1913) as This specimen is probably a duplicate of Ramalina combeoides Nyl. from Clifton­ the two given above for D. canescens.] by­the­Sea on dead Sambucus and *Ramalina farinacea (L.) Ach. — Bluff Isomera arborea. Cove: common on twigs of dead sage, *Niebla homalea (Ach.) Rundel & Hollinger 4386. See notes under R. Bowler — Clifton­by­the­Sea: on subleptocarpha below; Hasse may have limestone, 10 Sep. 1933, Darrow 314 collected this at Clifton­by­the­Sea, too. (ASU, WIS), as Ramalina homalea Ach., *Ramalina leptocarpha Tuck. — Bluff det. by P.A. Bowler in 2003. Cove: common on twigs of dead sage, Opegrapha brattiae Egea & Ertz — Hollinger 4387. Abalone Cove: on rocks on seashore, 24 *Ramalina pollinaria (Westr.) Ach. — Dec. 1952, Weber s.n. (COLO 77734). Bluff Cove: on twig of dead sage, Reported by Egea & Ertz (2007). Hollinger 4388. [Opegrapha varia Pers. — White Point: *Ramalina subleptocarpha Rundel & on twig, 1895, Hasse 386 (ASU, MIN); Bowler — Clifton­by­the­Sea: May 1908, on Rhus integrifolia, Dec. 1896, Hasse 14 Hasse 116 (ASU, MIN, SBBG). These (ASU, SBBG), as O. diaphora Ach. three specimens appear to be duplicates, (Hasse 1913). Possibly misidentified; the one at ASU determined by B.D. Ryan most specimens in the greater Sonoran in 1999 as R. farinacea, the one in SBBG Desert region were misidentifications for determined by H. Kashiwadani in 2002 as other species of Opegrapha (Ertz & Egea R. subleptocarpha, and the remaining one 2007).] at MIN is recorded as R. lacera (With.) *Opegrapha xerica Torrente & Egea — J.R. Laundon. Bluff Cove: on branch of Rhus at edge of *Rinodina gennarii Bagl. — Sacred beach, Hollinger 4404a. This was a thin, Cove: on rotten rock on ground in grove, tan, scurfy, lichenized crust, with Hollinger 4436c. epruinose, black lirellae, carbonized [Rinodina rinodinoides (Anzi) H. exciple closed below the apothecium, and Mayrhofer & Scheid. — Hasse (1913) hyaline hymenium turning red in Lugol's. mistakenly reported Buellia rinodinoides The spores were hyaline, 4­septate, ~20 x Anzi. on calcareous rocks at White Point 6­7 µm, ±constricted at septa, all lumina (Sheard 2004).] equal with rounded ends. *Roccella gracilis Bory — Clifton­by­the­ *Physcia adscendens (Fr.) H. Olivier — Sea: 19 Jan. and 1 Jul. 1914, Hasse 14779 Bluff Cove: on twigs of dead sage, (ASU), det. by A. Tehler in 2000 and 2001 Hollinger 4389. as R. difficilis Darb. and R. peruensis *Pyrrhospora quernea (Dickson) Körber Kremp. — Bluff Cove: on limb of Rhus; on twigs [Roccella montagnei Bel. — Clifton­by­ of dead sage, Hollinger 4393. the­Sea: Jun. 1914, Hasse s.n. (MIN 2960,

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664453). Probably misidentifications since *Verrucaria mimicrans Servít — Bluff this species is not known to occur north of Cove: on crumbling rock in gully, Mexico.] Hollinger 4418. This had an indistinct, *Sarcogyne plicata H. Magn. — Clifton­ chalky thallus, mostly immersed by­the­Sea: Nov. 1896, Hasse s.n. (NY perithecia, involucrellum that was thick 1221349), det. by K. Knudsen in 2011. For and black in upper third, uniformly thin, notes on this species and S. “privigna”, black exciple, and spores ~28 x 20 μm. see Knudsen & Kocourková 2011. [Verrucaria muralis Ach. — Clifton­by­ *Sarcogyne regularis — Bluff Cove: on the­Sea: 1912, Hasse s.n. (MIN 2484). crumbling rock in gully, Hollinger 4415. This is a plausible record, being frequent This calciphilous species is distinguished on calcareous substrates in California by pruinose black apothecia which turn (Knudsen & Kocourková 2012), but there reddish when wet, although note that is no dot for Palos Verdes on the range nonpruinose species of Sarcogyne can map in the Sonoran Flora (Breuss 2007), develop a thin pruina when growing on so it requires confirmation.] calcareous or sandstone substrates *Verrucaria nigroumbrina (A. Massal.) (Knudsen & Standley 2007). Servít — Portuguese Bend: 24 Dec. 1952, *Sigridea californica (Tuck.) Tehler — Weber s.n. (ASU), as V. nigrofusca. There Clifton­by­the­Sea: 1912, Hasse s.n. (MIN is a dot shown on the range map for V. 875434, 875436), as Schismatomma nigrofusca Servít in the Sonoran Flora californicum (Tuck.) Zahlbr.; Bluff Cove: suggesting that Breuss (2007) may have common on twigs of dead sage, Hollinger seen this specimen. 4412; Sacred Cove: on limb of dead *Verrucaria viridula (Schrader) Ach. — unknown shrub in grove, Hollinger 4429; Bluff Cove: on crumbling rock in gully, sheltered limb of Rhus on top of cliff, Hollinger 4419. This was a rimose to Hollinger 4432; limb of dead exotic shrub areolate, greenish to brownish crust, with at edge of cliff, Hollinger 4435. My perithecia immersed in the center of specimens determined by K. Knudsen. areoles except for slightly protruding black *Trapelia coarctata (Turner ex Sm. & tips, involucrellum restricted to the upper Sow.) M. Choisy — Clifton­by­the­Sea: portions and flaring away from exciple, on soil, Apr. 1914, Hasse 242 (ASU, exciple thick and hyaline, and spores ~25­ SBBG), as Lecidea coarctata (Turn. ex 30 x 15­20 μm. Sm.) Nyl. f. elachista