Ochrolechia raynori, a New Lichen Species from the Southern Rocky Mountains (Colorado, USA) and Key to Asexually Reproducing Ochrolechia in Western North America

Simple Summary

Around the world, lichens remain one of the least documented groups of macroogranisms. Many to most places worldwide have never had a formal lichen inventory, and this includes the southern Rocky Mountains of Colorado. Despite the incredible topographic relief and ecological heterogeneity present within Colorado’s borders, expected to harbor a uniquely rich and varied lichen biota, the lichens of the state have never been studied. Here, we describe a new, asexually reproducing lichen species from the cosmopolitan genus Ochrolechia from the Front Range Mountains of Colorado, and then provide a key to help identify all described asexually reproducing species of the genus in western North America. This new species is tentatively ranked as Critically Endangered per IUCN guidelines. Future fieldwork to inventory the lichens of Colorado by ourselves and colleagues, will, we predict, uncover new discoveries of additional populations of this species and other novelties.

Abstract

Ochrolechia is a diverse and charismatic lineage of both sexually and asexually reproducing lichens, with centers of species richness in northern temperate areas of the world, including North America. As part of recent work to comprehensively inventory the lichens of the Indian Peaks Wilderness (Arapaho–Roosevelt National Forest, Front Range Mountains, Colorado), we discovered material of a sorediate member of the genus to which no existing names could be applied. This material was collected in very remote, extremely difficult-to-access mid-montane forests of the west slope of the Indian Peaks Wilderness, in a steep and jagged off-trail drainage (Hell Canyon). Subsequent study of this material along with review of pre-existing collections at the COLO Herbarium revealed it to represent a new scientific species. We here formally describe Ochrolechia raynori, in honor of Seth Raynor who led the Indian Peaks Wilderness lichen inventory. We additionally document the occurrence of Dactylospora parasitica on this new lichen species. Ochrolechia raynori is distinctive for its continuous, smooth, shiny thallus that bears discrete soralia and coarse soredia, its occurrence on mosses and other lichens that overgrow rocks, and its chemistry. We generated a molecular phylogeny of this and other members of Ochrolechia using the nrITS locus and show O. raynori to be sister to the widespread, sexually reproducing species O. upsaliensis. This occurrence of an asexual species that is sister to a sexual species is consistent with the “species pair” hypothesis in lichenology, which suggests an intimate role of reproductive mode divergence in the process of speciation. Examination of the phylogeny yielded evidence of four additional pairs in Ochrolechia, for a total of five species pairs, which indicates that this phenomenon may be a common occurrence in this lineage. IUCN Conservation Assessment of Ochrolechia raynori revealed the species to be best considered as Critically Endangered. However, we expect that continued efforts to inventory the lichens of the southern Rocky Mountains, especially in some of its wildest, most remote regions in similar habitats, will likely result in the discovery of additional populations of this remarkable new species.

1. Introduction

Ochrolechia A. Massal. (Pertusariales M. Choisy ex D. Hawksw. & O.E. Erikss.) is a charismatic genus of crustose lichens consisting of >60 species worldwide, with centers of diversity including temperate regions of North America, Europe, and China [1,2,3,4]. The genus is characterized by a crustose thallus that is most commonly continuous to areolate, a trebouxioid photobiont (but see [5]), typically large and showy apothecia (when fertile), I+ amyloid asci (and hymenium), branched paraphyses, and simple, hyaline, typically large spores [6,7,8].

As currently understood, North America is home to ~38 species of Ochrolechia [1,6,9,10]. Of these, slightly over half reproduce primarily through sexual means (i.e., apothecia, meiotic ascospores) and the remainder, slightly under half, reproduce primarily through lichenized asexual reproductive propagules (e.g., soredia, isidia) which are products of mitosis and co-disperse the fungal, algal, and additional lichen constituents together ([11], Watts et al., ms in prep.). This remarkable relative balance of reproductive modes is unusual for many lineages of lichens, wherein sexual reproduction is much more of a dominant mode compared to asexual means (Watts et al., ms in review). Among these asexual species, chemistry is both diverse and taxonomically important, with frequently produced compounds including orcinol depsides, orcinol depsidones, β-orcinol depsides, fatty acids, and xanthones [3,6,12].

The common occurrence of both sexually and asexually reproducing species in Ochrolechia makes the lineage a natural system in which to test hypotheses of the evolution of “species pairs” ([11] and references therein), wherein sister species or closely related species are divergent in reproductive mode, with one member of the pair reproducing primarily through sexual means and the other reproducing primarily through asexual means. Such species pairs may in fact represent common occurrences in the genus, such as in the present new species and its closest relative.

The last taxonomic revision of Ochrolechia in North America was by Howard [1], followed by a revision of the corticolous species by Brodo [6]. In this study, we document and describe a new species of Ochrolechia to science from the southern Rocky Mountains of Colorado, then place this new diversity in the context of now 19 known asexually reproducing species in the genus in western North America through molecular phylogenetic analyses and a dichotomous key. This new species was discovered during one grueling field day by the first author together with Seth Raynor, who spearheaded an inventory of the lichens of the Indian Peaks Wilderness (Arapaho-Roosevelt National Forest, Front Range Mountains, Colorado; Raynor & Manzitto-Tripp, ms in revision). The asexually reproducing Ochrolechia raynori is here shown to be sister to the widespread, sexually reproducing species O. upsaliensis, and we hypothesize a “species pair” evolutionary divergence event between the two. We explore the remainder of the phylogeny for evidence of additional species pairs and find instances of four others, for a total of five species pairs. Finally, we document the occurrence of the lichenicolous species Dactylospora parasitica (Flörke) Arnold on this new species, representing its first known occurrence on the genus Ochrolechia and in the state of Colorado to our knowledge.

2. Material and Methods

Field Work. Throughout the summers of 2020–2023, the first author and second author joined colleague Seth Raynor for several field trips throughout the Indian Peaks Wilderness (IPW) of Arapaho–Roosevelt National Forest. This expansive and topographically complex region forms the basis for much of the skyline that lines the Colorado Northern Front Range, ranging in elevation from a low point of ~8400 ft. to a high point of ~13,500 ft. (North Arapaho Peak). During and after this time, collections to comprehensively document the lichen biota of Colorado have been made by all three individuals, throughout the state of Colorado, but with a second emphasis on the lichens of Boulder’s Open Space & Mountain Parks (Watts et al., ms in prep.).

Given a paucity of lichen collections from the west slope of the Continental Divide in the IPW, Raynor and the first author carried out one specific field trip on the west slope (Grand County) to inventory the middle montane habitat. We chose to target an extremely remote and rugged, difficult-to-navigate mesic canyon named Hell Canyon owing to its sheltered and shaded aspect. The canyon itself was so steep and narrow that the lowermost portions likely never receive direct sunlight, and were unlikely to have been surveyed by prior botanists. We sampled all lichen growth forms (i.e., foliose, fruticose, crustose, squamulose, leprose) and all substrates suitable for lichen growth (e.g., rock, bark, lignum, soil, bryophytes, sap, other lichens). Geographical locations of collections were recorded with an iPhone 14 Pro (Apple Inc., Cupertino, CA, USA), along with habitat characteristics. These locations were delimited opportunistically, i.e., based on habitat and biota that appeared interesting to the collectors. On the subsequent day, fieldwork, collections were pre-processed, i.e., stabilized (glued where necessary), then placed in temporary packets to prepare for final identifications (see below). Subsamples were removed for DNA sequencing then stored under refrigeration until returned to freezers in Manzitto-Tripp’s molecular laboratory at The University of Colorado Boulder.

2.1. Herbarium Study

Final identifications of all lichen collections were made at the University of Colorado, Museum of Natural History, COLO Herbarium [13] using an Olympus SZX16 and Olympus BX51 coupled to an Olympus EP50 camera. Samples were hand-sectioned then examined in water mounts along with other media. Spot tests using standard reagents (K, C, KC, P, N) along with UV illumination were conducted at COLO. Additionally, Thin Layer Chromatography and/or DNA sequencing was conducted by the authors at COLO (in Manzitto-Tripp’s molecular lab) on specimens to help confirm their identifications. Thin Layer Chromatography (TLC) utilized Solvent C and silica plates following Culberson & Kristinsson [14] and Culberson [15]. In addition to studying our own newly collected materials, we examined relevant material (i.e., all undetermined material already accessioned under Ochrolechia spp., n = 24 specimens) housed at COLO using similar methods to the above, providing updated identification annotations where necessary. All collections cited below were verified by the authors but, nonetheless, we include a “!” following any of the studied specimen citations, unless in reference to a type specimen housed elsewhere and only photographs were seen (in which case “photo!” is utilized). We referenced the identification of our collections using published literature, floras, field guides, existing herbarium material, and the Consortium of Lichen Herbaria (CLH, Lichen Portal; https://lichenportal.org/portal/, accessed 10 July 2025). Our taxonomy follows [9]. Data and specimens reported in this study are deposited at COLO and data were uploaded to COLO’s database, which is hosted on CLH.

2.2. Molecular Analyses

To place the new species in a molecular genetic context, we generated data from the nuclear ribosomal Internal Transcribed Spacer (nrITS) “barcoding” locus of the new species from the holotype (E. Manzitto-Tripp & S. Raynor 9795) and other members of the lineage (Pertusariales). Genomic DNA was extracted using a modified CTAB protocol [16]. We amplified the nrITS region (including 5.8 s) using ITS4 and ITS5 primers [17]. Resultant successful PCR products were sent to Quintara Biosciences (Cambridge, MA, USA) for product cleanup and bidirectional Sanger Sequencing. We generated a DNA alignment using our newly generated sequences together with pre-existing nrITS data retrieved from GenBank (GenBank numbers are provided in names of taxa/terminals in the phylogeny) and guided by existing literature, including outgroup selection. The final matrix consisted of 96 terminals.

Phylogenetic relationships within Pertusarales were constructed using Maximum Likelihood methods implemented in raxml 2.0 [18] and an ML + transfer bootstrap + consensus algorithm, with a GTR + FO + I + GA model of sequence evolution and 100 bootstrap replicates. We generated a 50% majority rule consensus tree from each matrix then visualized results using FigTree v1.4.4 [19]. We interpreted bootstrap support (BS) values indicative of support for a given phylogenetic relationship as follows: ≥90% = strong support, ≥80–89% = moderate support, ≥70–79% = weak support, and <70% = unsupported. Taxon identification, GenBank number, and collection locations for each specimen included in the phylogeny were displayed at branch tips.

2.3. Conservation Assessments

The conservation status of the new species was assessed following the International Union for Conservation of Nature (IUCN). Given a lack of useful historical data on population sizes and trends through time, which are necessary for criteria A, B, C, and E, we opt to utilize criterion D of the IUCN guidelines version 16 [20]. Under IUCN criterion D, species are classified as Critically Endangered (CR) or Endangered (E) if the total estimated population size is less than 50 individuals or between 51 and 250, respectively. We consider one mature individual as any 1 m² surface of rock containing an individual of the focal species, following practices recommended by a recent review about conducting conservation assessments of lichens [21].

3. Results and Discussion

That challenging day in Hell Canyon yielded material of a highly unique species of Ochrolechia that we were unable to identify using pre-existing names. As such, Ochrolechia raynori (Figure 1) is formally described below. The west slope of the IPW proved to be exceptionally rich (Figure 2), yielding at least one other new lichen species (Polycauliona pancakeana E. Tripp, Raynor, & J. Watts [22], the holotype of which derived approximately 0.5 miles from the Ochrolechia collection in the primary drainage into which Hell Canyon feeds (Buchanan Creek, Figure 2). Indeed, although we are in only the early phases of our long-term project to comprehensively inventory Colorado’s lichens, the initial discoveries of new species and new records have exceeded our expectations [22,23,24,25,26,27,28,29,30,31,32,33]. As such, it seems reasonable to hypothesize that a steep species accumulation curve will easily characterize the first 5–10 years of our work here.

Phylogenetic analyses resolved our new sorediate Ochrolechia raynori as sister to the widespread and sexually reproducing (“fertile”) species O. upsaliensis with strong support (100% BS; Figure 3). Indeed, O. raynori shares much of its secondary chemistry with O. upsaliensis, as discussed below (Part 1). The “species pair” sister relationship of these two taxa is similarly reflected in four other species-pair groups (for a total of five) reconstructed by our phylogenetic analysis: the isidiate O. yasudae as sister to the fertile O. xanthostoma (74% BS), the sorediate O. gowardii as sister to the fertile O. juvenalis (100% BS), the sorediate O. mahluensis as sister to the fertile O. frigida + O. tartarea. (99% BS), and the sorediate O. bahusiensis as sister to the fertile O. oregonensis (91% BS). Further phylogenetic analyses, with more complete sampling, will likely recover additional fertile-sterile species pairs in Ochrolechia. Wee recognize that the inference of sister relationships is highly susceptible to taxon sampling (hence, no formal character reconstruction analyses were conducted here given the limited nature of our sampling). Nonetheless, together, these five species pairs yield evidence supporting the hypothesis that reproductive mode transitions may be a common occurrence during speciation within the genus Ochrolechia (see [11] and references therein).

Combing of existing collections of Ochrolechia revealed only one additional collection of the species, which was made immediately adjacent to (north of) the IPW, i.e., Rocky Mountain National Park, but from the east slope (R. Anderson 3840 [COLO!], Boulder County; Figure 4). Additional fieldwork in similar habitats and at comparable elevations is needed to more fully reveal the range of this species. Following description and discussion of Ochrolechia raynori (Part 1), we present a key to identify asexually reproducing species of Ochrolechia in western North America (Part 2).

Part 1: TAXONOMY

Ochrolechia raynori E. Tripp & J. Watts, sp. nov.

TYPE: USA. Colorado. Grand County, Arapaho–Roosevelt National Forest, Indian Peaks Wilderness, Hell Canyon, extremely steep, mesic, and rocky slopes immediately bordering creek, with Acer glabrum, Alnus incana, Ceanothus velutinus, Picea engelmannii, Populus tremuloides, and Pseudotsuga menziesii, saxicolous and overgrowing mosses on rocks above creek, 9153 ft. elev., 40.11465–105.714750, 2 July 2023, E. Manzitto-Tripp & S. Raynor 9795 (Holotype: COLO!; Isotype: KANU!).

Mycobank #: 859527

Description. Thallus crustose, continuous, moderately thick, smooth except for swelling in the vicinity of developing soralia or as a function of underlying substrate (e.g., verruculose where underlying moss leaves depart from main axis), shiny, continuously corticate, whitish gray to faintly bluish gray. Prothallus absent. Upper cortex of two distinct layers, epinecral layer up to 35 µm thick, the eu-cortex paraplectenchymatous and composed of irregularly isodiametrical cells, up to 25 µm thick. Algal layer continuous, 35–75 µm thick. Medulla well-developed, up to 175 µm thick directly underneath a developing soralium or up to 50 µm thick when not associated with a soralium. Soralia round and regular in shape, discrete, sometimes coalescing with age but always remaining conspicuously distinct from one another, never entirely covering the upper surface. Soredia light grayish white, almost always slightly lighter in color than that of the thallus, 63–125 µm in diameter. Apothecia and associated sexual reproductive structures not seen.

Etymology. Ochrolechia raynori is named in honor of our dear colleague Seth Raynor, graduate student at the University of Colorado (EBIO Department), who spearheaded an impressive inventory of the lichens of the Indian Peaks Wilderness where this new species was discovered. The species was found in Hell Canyon, a remote and difficult-to-traverse drainage on the west slope of the range, where both the primary collector and co-collector (Raynor) endured considerable hardship during fieldwork. Seth’s unwavering dedication to understanding the lichen biota of Colorado combined with his resilience and remarkable work ethic in the field and herbarium have led to significant contributions to the study of lichens in the region. His ongoing research continues to expand and enrich our knowledge of Colorado’s diverse lichen flora. We are deeply grateful for Seth’s tireless efforts, collaboration, and most excellent companionship during our journey to understand these fascinating organisms in the southern Rocky Mountains.

Chemistry. Spot Tests: K-, C-, KC-, P-, UV- (thallus and soredia). Thin Layer Chromatography: murolic acid complex unknown (charring brown, below gyrophoric, above variolaric), unknown fatty acid (just above or level with the previous chemical), variolaric acid.

Distribution. Ochrolechia raynori, as currently understood, has a highly restricted range and is so far known only from the west slope (Grand County) and east slope (Boulder County) of the Front Range Mountains of Colorado. The first derived from the Indian Peaks Wilderness of Arapaho–Roosevelt National Forest and the second derived from Rocky Mountain National Park, immediately north of and adjacent to the IPW.

Substrate and Habitat. Ochrolechia raynori has been found overgrowing mosses and lichens on rocks on the banks immediately above Hell Canyon Creek in a mesic upper montane forest. On the east slope of the continental divide in Wild Basin, it was also found overgrowing saxicolous lichens in a mesic upper montane forest, suggesting a narrow niche.

Notes. Ochrolechia raynori is distinguished by its shiny, continuous, smooth thallus, discrete (almost pustulose) soralia that bear very coarse soredia, occurrence overgrowing mosses and lichens on rocks, and chemistry. Chemically, it is the only species with an unknown fatty acid plus an unknown compound in the murolic acid complex and variolaric acid, the latter two compounds shared with Colorado material of Ochrolechia upsaliensis (e.g., E. Tripp 5112 [COLO]), to which the new species was resolved as sister, phylogenetically.

Of sterile species of Ochrolechia in North America that produce variolaric acid, Ochrolechia raynori is unique for its production of secondary metabolites in the murolic acid complex, its negative spot tests, and its occurrence on moss and detritus rather than bark and wood, which is more common for species in this group. In general, TLC is needed for reliable identification of species of Ochrolechia containing variolaric acid. Perhaps most similar to the new species is Ochrolechia alboflavescens (Wulfen) Zahlbr.; however, the latter produces lichesterinic and protolichesterinic acids rather than murolic acid complex compounds. Ochrolechia farinacea Howard similarly produces murolic acid complex compounds, but occurs on bark, especially that of Quercus, has coalescing pale yellow soredia, and produces abundant UV+ bright yellow apothecia. Ochrolechia turneri (Sm.) Hasselrot differs by its occurrence on bark and lack of the murolic acid complex. Ochrolechia microstictoides Räsänen is known only from maritime New England and Nova Scotia in North America and differs in its production of lichesterinic acid, which results in a UV+ white reaction of the thallus. Ochrolechia gowardii Brodo produces variolaric acid and gyrophoric acid, yielding a C+ reaction in the soralia. Two additional species fail to produce either variolaric or gyrophoric acid: Ochrolechia sorediosa Howard differs by its occurrence on bark and its diffuse, coalescing soralia, that are P+ yellow. Ochrolechia minuta (Degel.) T. Sprib. occurs on bark and produces alectoronic acid, reacting KC+ pale red and UV+ white. We provide a key to sterile Ochrolechia in western North America to aid in the identification of the new species with respect to relatives. For more detailed information and useful taxonomic treatments of Ochrolechia in North America, see Howard [1] for a revision of all species, and Brodo [6] for a treatment of corticolous species and more detailed chemistry.

The holotype of Ochrolechia raynori (E. Manzitto-Tripp & S. Raynor 9795) is furthermore noteworthy for its lichenicolous ascomycete Dactylospora parasitica (Flörke ex Sprengel) Zopf. So far, this has not yet been documented to occur on Ochrolechia but rather is known from species of Pertusaria (Pertusariales). This is additionally the first documented occurrence of Dactylospora parasitica in Colorado.

In most clades of lichenized fungi, fertile species appear to never produce asexual lichenized propagules (in contrast, asexual species can sometimes manufacture sexual, fruiting bodies). However, in Ochrolechia, workers have noted the apparently increased tendency for fertile species to occasionally produce soredia [3,6]. The phenomenon is interesting and warrants further study, as it may ultimately impact stable taxonomy in Ochrolechia. In the case of O. raynori, however, we are unaware of any reports of sorediate in O. upsaliensis, despite it being common and widespread throughout the arctic. Additionally, the holotype of O. raynori is wholly sorediate and bears no fruiting bodies. We therefore here treat O. raynori at the species level rather than a sorediate form of O. upsaliensis.

Conservation Assessment. Ochrolechia raynori is here assessed as Critically Endangered (CR) under IUCN criterion D because of its small population size (two mature individuals currently known) and its highly restricted niche of near-pristine, riparian, mesic, upper montane forests of the northern Front Range Mountains. We note, however, that this assessment should be considered provisional given that much of the landscape remains unexplored from a lichenological perspective. New fieldwork (along with searches of relevant herbarium holdings), especially work that targets suitable habitat, may recover additional known populations of the species.

Additional Specimens Examined. USA. Colorado. Boulder County. Rocky Mountain National Park, Wild Basin, southwest of campground, 8600–8800 ft. elevation, 26 April 1963, R. Anderson 3840 (COLO!).

PART 2: Key to sterile Ochrolechia in western North America (north of Mexico)
1a.	Thallus isidiate or producing isidia-like spines	2
1b.	Thallus sorediate, not producing isidia-like spines	6
2a.	True isidia absent, isidia-like spines present, soredia sometimes present, usually
	terricolous, arctic-boreal distribution	3
2b.	True isidia present, isidia-like spines absent, soredia sometimes present, substrate
	and distribution various	4
3a.	Upper portions of cortex C+ yellow, containing variolaric acid, lower portions of
	cortex C+ red, containing gyrophoric acid	Ochrolechia alaskana (Verseghy) Kukwa
3b.	Entire cortex C+ red, containing gyrophoric acid	Ochrolechia frigida f. lapuensis (Vain.) Coppins
4a.	Isidia soon breaking down into granules to form a granular crust, in arid habitats,
	usually on Juniper wood	Ochrolechia subisidiata Brodo
4b.	Isidia not breaking down into granules, in humid maritime habitats, on wood 5
5a.	Isidia densely branched, occurring in clusters, usually growing on conifers, known
	from coastal southern Alaska	Ochrolechia cooperi T. Sprib.
5b.	Isidia thick, cylindrical, occasionally branched, occurring uniformly across the thallus,
	growing on hardwoods, distribution primarily eastern, but also reported
	from coastal Washington and Oregon	Ochrolechia yasudae Vain.
6a.	Cortex, medulla, and/or soralia C+ red, KC+ red, containing gyrophoric and/or
	lecanoric acids	7
6b.	Cortex, medulla, and/or soralia C+ yellow or C−, KC− or KC+ yellow, or KC+
	fleeting pink/purple, containing variolaric acid, alectoronic acid, or unknown
	substances	13
7a.	Soralia (and cortex) UV+ vibrant yellow-orange, containing lichexanthone	Ochrolechia arborea (Kreyer) Almb.
7b.	Soralia UV− or UV+ white, lacking lichexanthone	8
8a.	Cortex C−, soralia and epihymenium C+ red	Ochrolechia gowardii Brodo
8b.	Cortex C+ red, containing gyrophoric and/or lecanoric acids, soralia and
	epihymenium spot test various	9
9a.	Cortex K+ yellow, containing atranorin, always terricolous, distribution
	arctic-boreal	10
9b.	Cortex K−, lacking atranorin, usually corticolous or lignicolous, distribution
	various	11
10a.	Thallus thick, well developed	Ochrolechia gyalectina (Nyl.) Zahlbr.
10b.	Thallus thin, poorly developed	Ochrolechia inaequatula (Nyl.) Zahlbr.
11a.	Thallus thin, rimose-areolate, lacking accessory compounds by TLC	Ochrolechia mahluensis Räsänen
11b.	Thallus thin or thick, containing accessory compounds (other than gyrophoric and
	lecanoric acids) by TLC	12
12a.	Thallus usually thin, containing fatty acids of the murolic acid complex, lacking
	hiascic acid	Ochrolechia bahusiensis H. Magn.
12b.	Thallus usually thick and bullate-verrucose, lacking murolic acid complex fatty acids,
	containing other unknown fatty acids and sometimes hiascic acid	Ochrolechia androgyna (Hoffm.) Arnold
13a.	Cortex C+ yellow	14
13b.	Cortex C−	15
14a.	Apothecial margins and disks UV−, soralia spot tests variable, containing variolaric
	acid in low concentrations	Ochrolechia alboflavescens (Wulfen) Zahlbr.
14b.	Apothecial margins and disks UV+ yellow orange, soralia C+ yellow, containing an
	unknown UV flourescent compound and variolaric acid in high
	concentrations	Ochrolechia farinacea Howard
15a.	On mosses and detritus over rock, containing unknown compounds of the murolic
	acid complex	Ochrolechia raynori E. Tripp & J. Watts
15b.	On bark, lacking murolic acid complex compounds	16
16a.	Soralia C+ yellow, UV+ glaucus orange or UV−, P−	Ochrolechia turneri (Sm.) Hasselrot
16b.	Soralia C−, UV+ bright white or UV−, P+ pale yellow or P−	17
17a.	Soralia UV+ bright white, P−, containing alectoronic acid, distribution
	coastal	Ochrolechia minuta (Degel.) T. Sprib.
17b.	Soralia UV−, P+ pale yellow, containing an unknown compound, distribution
	inland	Ochrolechia sorediosa Howard