Open Access
1 March 2016 A Review of the Lichens of the Dare Regional Biodiversity Hotspot in the Mid-Atlantic Coastal Plain of North Carolina, Eastern North America
James C. Lendemer, Richard C. Harris, Ana Maria Ruiz
Author Affiliations +

The results of a large-scale biodiversity inventory of lichens (including lichenicolous and allied fungi) in the Dare Regional Biodiversity Hotspot (DRBH) are presented. The DRBH is a region within the Mid-Atlantic Coastal Plain (MACP) of eastern North America that was recently delineated based on its unique and diverse lichen communities relative to other areas of the Atlantic Coast. Drawing on 4,952 newly generated voucher specimens from 49 sites, patterns of biodiversity and biogeography are presented and discussed within the context of both the DRBH and the broader MACP. Relationships between natural communities, vegetation, and lichen communities are discussed, as are threats to the lichen biota. A series of conservation actions are presented together with avenues for future study. In addition, supplementary resources are provided in the form of: (a) a checklist of DRBH lichens, lichenicolous fungi, and allied fungi; (b) keys to DRBH lichens and lichenicolous and allied fungi; and (c) formal descriptions of the following species new to science that were discovered during the inventory: Albemarlea pamlicoensis gen. et. sp. nov., Arthonia agelastica sp. nov. (on Lecanora louisianae B. de Lesd.), Arthonia hodgesii sp. nov. (on Graphis lineola), Arthonia stevensoniana sp. nov. (on Haematomma accolens), Lichenochora haematommatum sp. nov. (on Haematomma persoonii), Megalaria alligatorensis sp. nov., Minutoexcipula miniatoexcipula sp. nov. (on Pertusaria epixantha), Trichosphaerella buckii sp. nov. (on Punctelia rudecta).

INTRODUCTION In 2012, we began a large-scale and systematic inventory of lichen biodiversity in the Mid-Atlantic Coastal Plain (MACP) of eastern North America. The MACP is an ecoregion approximately 89,691 km2 (34,630 mi2) in size that comprises the low-lying ecosystems along the Atlantic Coast of the USA from southern New Jersey to northern Florida (US Environmental Protection Agency 1997, Auch 2000). The MACP is recognized as one of the most biologically diverse ecoregions in North America (Hall and Schafale 1999), although its lichen biota had been the subject of little study. Overall, the region has been highly impacted by anthropogenic change (Auch 2000, Loveland and Acevedo 2000, Napton et al. 2010), with nearly a tenth of the region remaining as “intact natural habitat,” which is increasingly threatened by diverse forces, including fragmentation and loss of remaining habitat by development, increased storm intensity and frequency, overexploitation of timber resources, pollution, and sea level rise (Kirby-Smith and Barber 1979, LeGrande et al. 1992, Bellis 1995, Shankman 1996, Riggs and Ames 2003, Brown et al. 2005, Berman and Berquist 2007, Hupp et al. 2009, Sallenger et al. 2012, Villarini and Vecchi 2012, Lorber and Rose 2015, US Army Corps of Engineers 2015).

Before beginning our inventory of the MACP, we expected, based on experience from previous small-scale efforts in the region, that we would encounter considerable diversity as well as many new and unusual species. However, when we began to inventory the largest remaining contiguous protected areas in the MACP, an extensive area of swamps centered in the Albemarle-Pamlico Peninsula of North Carolina, we immediately recognized that the region hosted much higher levels of lichen biodiversity than we had expected. Following two years of field study, we summarized our biodiversity data for the portion of the MACP between southern New Jersey and the North Carolina–South Carolina border. This led to the discovery that all of the most biodiverse sites in that portion of the MACP were concentrated on or near the Albemarle-Pamlico Peninsula, and that they were all within 1 m of current sea levels (Lendemer and Allen 2014). This area largely corresponds to the original extent of the once vast swamp that was historically referred to as the Great Alligator Dismal (e.g., Morse 1804, Cummings 1966). In recognition of the unique lichen communities in the Albemarle-Pamlico Peninsula, we designated the region as the Dare Regional Biodiversity Hotspot (DRBH), a lichen biodiversity reservoir imperiled by sea-level rise and anthropogenic change (Lendemer and Allen 2014).

Recognizing the importance of a thourough understanding of the lichens of the DRBH for the effective conservation of American, let alone global, lichen biodiversity, we held the 23rd Tuckerman Workshop in the region in 2014. Before and during the workshop, we visited additional localities, and considerably increased our knowledge of the known biodiversity. The remarkable increase in DRBH lichen biodiversity found as a result of the Tuckerman meeting convinced us that a summary treatment was needed, and that such a work would be useful to both the lichenological community and, more broadly, to those involved in the biodiversity sciences, as well as conservation and management in the MACP. As such, we undertook the present study, the results of which are presented here.


Delineation of the Study Area

This study details the results of a large-scale biodiversity inventory of the DRBH (Figure 1A), an area recently designated by Lendemer and Allen (2014) because of its unique and outstandingly diverse lichen communities. The DRBH is located entirely within the North Carolina portion of the EPA Level III Ecoregion the MACP (US Environmental Protection Agency 1997, Auch 2000). The bulk of the DRBH is comprised of the mainland Albemarle-Pamlico Peninsula, a peninsula bounded to the north by the Albemarle Sound and to the south by the Pamlico Sound, together with the adjacent barrier islands that form a portion of the famous North Carolina Outer Banks. As delineated here, the DRBH also includes the North River drainage, which is an extensive system of swamp forests along the North River just north of the Albemarle-Pamlico Peninsula in mainland Camden and Currituck Counties. Essentially, the DRBH comprises all of the North Carolina counties of Currituck, Dare, Hyde, Tyrrell, and Washington, together with a portion of Camden County located along the North River (Figure 1B). This area encompasses the largest contiguous natural areas in the MACP (Lendemer and Allen 2014) and has a land area 4,501 km2 in size, of which 1,604 km2 (36%) are protected.

Figure 1. 

A. Satellite image maps illustrating the study area and surrounding region. B. The counties that make up the study area. C. All georeferenced vouchers available in Consortium of North American Lichen Herbaria as of 2015 that were collected prior to the present study. D. The sites inventoried as part of this study.


Field Inventory and Herbarium Study

This study is based on a combination of fieldwork completed as part of a lichen biodiversity inventory of the MACP (see Lendemer and Allen 2014), and a large-scale study of existing vouchers deposited in the herbarium of the New York Botanical Garden (NY). Field work was carried out over a series of four trips, beginning with 8–12 December 2012 carried out by J.C.L., R.C.H., and W.R. Buck, followed by 18–19 March 2013 by J.C.L., 23–24 March 2013 by J.C.L., J.L. Allen, and E.A. Tripp, and 18–24 March 2014 by 30+ participants of the Tuckerman Workshop, including all of the individuals of the previous trips. The Tuckerman Workshop is an annual meeting wherein professional, amateur, and student lichenologists spend five days of field and laboratory time inventorying lichen biodiversity of a region of eastern North America as a group.

Field methods followed those outlined by Lendemer and Allen (2014), wherein a team spent 1–2 hr conducting an expert-based inventory using floristic habitat sampling (see Newmaster et al. 2005) within a site (3.43 ± 2.2 ha) delineated in such a way as to be uniform in habitat (i.e., a single vegetation type with uniform elevation). Sites were selected spontaneously in the field based on direct observations of habitat quality and lichen diversity from vehicles during visits to as many protected areas as were allowed by available time and funds. This method of site selection was intentionally nonrandom, because the goal of the inventory was to document as much biodiversity as possible given available time and resources. Thus, the lowest-diversity sites and most degraded habitats were excluded from this study based on the extensive field experience of the two senior authors in conducting large-scale biodiversity inventories.

Teams of approximately equal numbers allocated approximately equal time to inventory across sites. Each member of the team collected a voucher of each taxon that they encountered at each site, with the goals of: (a) comparing the taxonomic overlap between collectors; (b) including all common species; and (c) capturing the most complete substrate diversity possible for each species. For each voucher, the substrate and microhabitat were recorded in the field, and for each site, the overall vegetation, habitat, and condition of the site were characterized by field observation.

All vouchers were identified using existing published (e.g., Harris 1995) and unpublished (e.g., the keys published herein) resources. Specimens were examined dry using an Olympus SZ-STB dissecting microscope (Olympus). Thallus anatomy and measurements of microscopic characters were carried out on sections prepared by hand with a razor and mounted in water, viewed using an Olympus BX53 compound microscope equipped with a DP72 digital camera (Olympus) and CellSens imaging software (Olympus). Chemistry was studied with standard spot tests (K, C, P, UV) following Brodo et al. (2001), and with thin-layer chromatography (TLC) using solvents A and C following Culberson and Kristinsson (1970), as modified for the peanut butter jar by Lendemer (2011).

After identification, all specimens were labeled with georeferenced locality data, voucher-specific substrate data, and collection information. They were then digitized in KEMu and assigned unique identifiers in the form of barcodes and electronic record numbers. Data digitization was performed by one individual (J.C.L.) to assure that records were entered uniformly, taxonomic identifiers were kept constant, and that variants of georeferenced locality data were not entered multiple times. During incorporation into the NY herbarium, any existing vouchers from the MACP already in the collection were also examined, the identifications verified, localities georeferenced, and the data digitized. A special effort was also made to locate, identify, curate, and digitize any additional MACP vouchers located in the undetermined material at NY, as well as in unprocessed portions of donated herbaria, including those of J.P. Dey, E. Lay, and C.F. Reed.

Dataset Assembly, Analysis, and Visualization

Data used in analyses were exported from the NY KEMu system as a single CSV file containing all records from North Carolina. A master data file was created from this bulk export by pruning the dataset to include only records of lichens and lichenicolous fungi from MACP counties following US Environmental Protection Agency (1997). The master dataset was then copied and manipulated as follows: (a) individual CSV files were created for each species for use in producing species distribution maps; and (b) the master file was pruned to include only the subset of records from the DRBH. The pruned dataset composed only of DRBH records was then used to generate a taxonomic checklist for the DRBH, and to obtain counts for the number of vouchers (unique herbarium specimens) and occurrences (locations defined as a unique latitude and longitude point) for each taxon. The DRBH dataset was then further pruned to include only records from sites inventoried by our team as part of the four trips outlined above. This smaller dataset was then used to: (a) obtain taxonomic diversity values (total number of unique taxonomic identifications) for each site; (b) generate a species versus site presence/absence matrix; and (c) generate a species × collector × site presence/absence matrix for the subset of sites visited by three team members (J.C.L., R.C.H., and W.R. Buck).

An additional dataset comprising all records from the DRBH available in the Consortium of North American Lichen Herbaria (CNALH) online database was downloaded on 1 June 2015. Nomenclature for the dataset was updated following Esslinger (2014), and the dataset was pruned to contain only prestudy DRBH voucher data as follows: (a) records erroneously included that were not actually from the DRBH were removed; (b) records of vouchers generated as part of this study or by participants during study-related workshops (e.g., the Tuckerman Workshop held in the DRBH in 2014) were removed.

ArcMap 10.0 (ESRI 2011) software was used to plot the coordinates and geospatially visualize data for the purposes of this study. ArcGIS World Imagery baselayer (ArcGIS World Imagery, Redlands, CA) was selected for the map background, with features including 0.3-m resolution imagery in the continental USA. Georeferenced species occurrence data were saved in CSV-formatted files and uploaded to ArcMap. For each set of occurrence data or site coordinates, the geographic coordinate system for the dataset was selected to display the World Geodetic System 1984. The dataset and map layers were then exported and saved as a single shapefile document. Through ArcToolbox: Data Management Tools, the shapefile was projected using North American Lambert Conformal Conic as the output coordinate system. Routine data calculations were performed, summarized, and visualized in Microsoft Excel (Microsoft Inc., Redmond, Washington). Similarity values and other diversity statistics were calculated using EstimateS for Windows (v9.10; see Colwell and Elsensohn 2014) from datasets formatted as tab-delimited TXT files. All datasets used in this study are archived in Dryad as doi  10.5061/dryad.226d0.

RESULTS AND DISCUSSION A total of 49 sites within the DRBH were inventoried for this study (Figure 1D), from which 4,952 voucher specimens were collected, representing 386 taxa. Of these 386 taxa, 8 are described as new to science (Appendix I), and several were previously described as new to science as part of our inventory efforts (Lendemer and Harris 2014a, Lendemer and Goffinet 2015, Lendemer and Harris 2015). An additional 76 pre-existing vouchers from the DRBH were located at NY and also included in the study. The small number of existing vouchers reflects the overall lack of study that much of the MACP had received previously. This is further evidenced by the lack of mainland DRBH collections in the CNALH, where the majority of existing vouchers from outside this project were collected from coastal barrier islands (Figure 1C; and see “Comparison with Existing Data” section below).

A checklist of the lichens and lichenicolous and allied fungi of the DRBH is provided here in Appendix II. To facilitate further study of the DRBH lichen biota, and to improve the usefulness of this contribution, identification keys for the DRBH lichen biota are also provided here in Appendix III. These resources add to the small number of lichen floristic contributions for the MACP (Lendemer and Yahr 2004, Lendemer and Knapp 2007, Hodkinson and Case 2008), and are the first comprehensive taxonomic keys to cover any part of the Coastal Plain of southeastern North America. While an excellent floristic account, including keys, has been published for part of the Coastal Plain in northeastern North America (Brodo 1968), that work did not include the vast majority of the species present in the southeastern Coastal Plain, and is now taxonomically outdated. The results and discussion presented below are grouped by topic and intended to summarize the data generated from our inventory, place them within the context of the DRBH as well as other inventories, and further place the lichen biota of DRBH within the broader context of the MACP and the Atlantic Coastal Plain as a whole.

Overview: Vegetation and Lichen Diversity

The unexpected diversity of lichens and lichen communities found in the DRBH is an excellent illustration of the issues that have traditionally led to the Coastal Plain having been overlooked as a biodiversity hotspot (Noss et al. 2015). In our case, lichen biodiversity is concentrated in low-lying swamp forests and upland hardwood forests, in contrast to the longleaf pine savannas and wetlands that have been the focus of previous biodiversity conservation efforts in the region (see below). Even after having conducted extensive field studies in the region, it remains surprising to us that so many ecosystems, vegetation types, and lichen species could occur in an area with little topographic relief, and especially one that has been so greatly impacted by centuries of anthropogenic change (LeGrande et al. 1992, US Environmental Protection Agency 1997, Hall and Schafale 1999, Ricketts et al. 1999, Brown et al. 2005). That such heterogeneity and diversity exists in the DRBH, and more broadly within the Coastal Plain, is attributable to a suite of abiotic factors and stochastic events that have resulted in very different environmental conditions (e.g., soil types, hydrological regimes, microclimates) in close proximity and often at small scales (US Environmental Protection Agency 1997). In the case of the DRBH, what at first glance may appear to be a vast monotonous swamp is, upon closer examination, a rich mosaic of varied natural communities at both large and small scales (Lynch and Peacock 1982a, 1982b; LeGrande et al. 1992, US Department of the Interior 2007, 2008, Sorrie 2014a, 2014b).

From the lichen perspective, habitats in the DRBH can be classified into four main types: marshes and other nonforested wetlands; forested wetlands (i.e., swamps); peatlands (i.e., pocosins and Atlantic white cedar forests); and forested uplands. For an excellent summary of the natural communities within these groups, the reader should refer to Schafale and Weakley (1990); however, it should be noted that our four primary habitat types do not correspond directly to those outlined in that work. Rather, we have grouped the narrowly defined natural communities of Schafale and Weakley (1990) into broad groups (e.g., their eight pocosin and peatland communities are here treated collectively as “peatlands” with two main types). Refer to Table 1 for a comparison of the DRBH natural communities recognized here and those defined by Schafale and Weakley (1990).

Table 1. 

Comparison of Dare Regional Biodiversity Hotspot natural communities defined by Schafale and Weakley (1990) to those discussed in the present study.


Marshes and other nonforested wetlands are biologically productive, important habitats and are the focus of considerable study and conservation action (Odum et al. 1984, Benoit and Askins 2002, Kushlan et al. 2002, Street et al. 2004). Nonetheless, based on field observations in this study, these habitats support almost no lichen diversity, and thus were not inventoried (Lendemer and Harris, pers. obs.). This paucity of lichens in nonforested wetlands is due to the absence of suitable substrates, namely rocks, robust woody vegetation, and organic matter not submerged by saltwater.

Of the three types of forested habitats in the DRBH, the region is dominated by low-lying forested wetlands and slightly elevated peatlands. In fact, within the two largest protected areas of the DRBH, Alligator River and Pocosin Lakes National Wildlife Refuges, more than 232 km2 and 615 km2 of land fall into these groups, respectively (US Department of the Interior 2007, 2008). The taxonomic diversity of lichens found in forested wetlands, peatlands and uplands is summarized and compared in Table 2.

Table 2. 

Summary of lichen diversity across Dare Regional Biodiversity Hotspot (DRBH) habitat types as well as the pairwise similarities of their overall lichen communities.


By far, peatlands comprise the largest natural communities in the DRBH in terms of total land area, and these are dominated by different types of pocosins (see Figure 2 for examples). Due to the dense vegetation and frequent deep canals surrounding them, pocosins are difficult to access, let alone inventory. During our work in the MACP, we found only one species that appeared to be restricted to pocosins, a new species of Megalaria described herein, which was found twice at one site in Alligator River National Wildlife Refuge. Otherwise, the lichen communities of pocosins are composed of species that are widespread in adjacent swamp forests where understories host low lichen diversity, because they are extremely shaded. Nonetheless, future studies of pocosins, particularly focusing on the upper boles and canopy, could reveal additional specialized taxa not collected during our inventory.

Figure 2. 

Pocosin habitats in the Dare Regional Biodiversity Hotspot (all from Dare County). A. Marshland grading to pond pine (Pinus serotina)–shrub pocosin. B. Pond pine–shrub pocosin. C. Pond pine–cane (Arundinaria) pocosin with loblolly bay (Gordonia lasianthus). D. High shrub pocosin dominated by shrubs and Smilax with sparse pond pine.


The nonpocosin peatlands in the DRBH are composed of Atlantic white cedar (Chamaecyparis thyoides (L.) Britton) forests (Figure 3A). Atlantic white cedar typically forms dense, even-aged stands and is dependent on specific conditions for regeneration (Laderman 1989). The species has been extensively logged in the past, and the peatlands it forms ditched and drained, such that the ecosystem is now considered globally threatened (Laderman 1989, Burke and Sheridan 2005). Our inventory of Atlantic white cedar peatlands in the DRBH was limited primarily because only a small number of stands were deemed suitable for inventory. The majority of candidate sites were ruled out based on field observations that they hosted few if any lichens, owing to young stand age wherein tree stems were densely crowded and small. Furthermore, many of the remaining stands were impossible to access without specialized equipment or logistical support. We did inventory one pure Atlantic white cedar stand, as well as several stands where the species was present as a minor component of the forest. Our inventory largely confirmed prior studies of Atlantic white cedar peatlands (Torrey 1933, Thomson 1935, Little 1951, Brodo 1968, Harris 1985, Lendemer 2006; Lendemer unpubl. data from Delmarva and New Jersey) that have found that, although the habitat has a distinctive community of lichens, the same species also occur in swamp forests on other coniferous hosts, particularly cypress and pine. This phenomenon is illustrated by species such as Chrysothrix chamaecyparicola and Micarea chlorosticta, which often dominate corticolous Atlantic white cedar communities in North Carolina and elsewhere, but also occur less frequently and less abundantly on cypress and pine in other swamp forests (Lendemer and Elix 2010; Lendemer, unpubl. data).

Figure 3. 

Swamp forest habitats in the Dare Regional Biodiversity Hotspot. A. Atlantic white cedar (Chamaecyparis thyoides) swamp in Dare County. B. Bald cypress (Taxodium distichum) swamp in Tyrrell County. C. Tupelo (Nyssa) swamp in Dare County. D. Red maple (Acer rubrum) –dominated mixed hardwood swamp in Tyrrell County.


The most diverse lichen communities in the DRBH occur in the swamp forests, which, unlike peatlands, usually have relatively widely spaced trees and comparatively open understories (Figures 3B–3D; Kellison and Young 1997, Lorber and Rose 2015). Presumably, the abundance of diverse microhabitats stemming from more extreme light and humidity gradients, regularly fluctuating water levels, and higher diversity of hosts (i.e., many different hardwoods and conifers, often in varying composition) are factors that have fostered rich and diverse lichen communities in the swamp forests of the DRBH. However, it is significant that the DRBH also includes some of the largest and most intact stands of mature swamp forests that we encountered in the MACP (see, e.g, Sorrie 2014a, Lorber and Rose 2015). The low degree of past disturbance and larger contiguous forested areas relative to other parts of the MACP may also have facilitated the survival of lichens in the DRBH that were once more common and widespread in the Coastal Plain. This is evidenced by the finding that the DRBH, and specifically mature swamp forests, host the highest cyanolichen diversity of any sites we inventoried in the MACP, including species that we did not encounter elsewhere, such as Parmeliella pannosa (Sw.) Müll. Arg. and Pannaria tavaresii P.M. Jørg. (Lendemer and Goffinet 2015). Furthermore, the area with the richest cyanolichen communities in the DRBH is also where we discovered a new species of the macrolichen genus Sticta, S. deyana Lendemer & Goffinet (see Lendemer and Goffinet 2015). That species is locally common in a small area of Alligator River National Wildlife Refuge, but otherwise known from a single mature hardwood forest in Alabama. Sticta is an easily recognized and conspicuous genus, and the distribution of S. deyana almost certainty reflects actual rarity, rather than a lack of adequate lichen exploration in southeastern North America.

Uplands in the DRBH fall into two categories: protected maritime forests on barrier islands (Figures 4B–4D) and small isolated inland “islands” of upland surrounded by swamp forests (Figure 4A). Historically, there would have been large areas of upland forests on the inner portions of the DRBH mainland; however, these have been almost entirely converted for sylviculture, agriculture, and other uses (Dey et al. 2010, Noss et al. 2015). The small number of remaining upland forest sites, most of which are confined to barrier islands, limited our ability to inventory upland forests as extensively as the much larger and more intact swamp forests. Even the remaining upland forests on barrier islands have been severely fragmented and degraded (Lopazanski 1987, Bellis 1995, Berman and Berquist 2007). Nonetheless, maritime forests were the second most diverse habitat inventoried, with 207 taxa or 54% of the total known from the DRBH. Although these habitats share many species with inland swamp forests, they also have a distinct group of species that does not occur on the mainland, and even includes narrow endemics, such as Phaeographis oricola Lendemer & R.C. Harris (Lendemer and Harris 2014a, Lendemer and Harris 2015).

Figure 4. 

Upland habitats in the Dare Regional Biodiversity Hotspot. A. Mixed hardwood forest dominated by oaks (Quercus) and maple (Acer) with dense understory of cane (Arundinaria), Tyrrell County. B. Maritime forest dominated by loblolly pine (Pinus taeda) with an understory of mixed shrubs dominated by yaupon (Ilex vomitoria), Dare County. C. Stabilized dune scrub dominated by lichen ground cover (Cladonia evansii, C. leporina, C. subtenuis) and live oaks (Quercus virginana), Dare County. D. Maritime forest dominated by live oak, Currituck County.


Although we inventoried only one inland upland forest in the DRBH, that site hosted 20% (77 taxa) of the total DRBH diversity, and was unlike all other sites that we inventoried. The site consisted of a narrow upland ridge surrounded by a mixed hardwood swamp populated with relatively mature beech (Fagus grandifolia Ehrh.) forest. The recently described species Acanthothecis paucispora Lendemer & R.C. Harris was located at this site, and remains known from only two strongly disjunct locations that are both in the MACP (Lendemer and Harris 2014a). A crustose lichen that produces conspicuous pycnidia and apothecia, with polysporous asci containing numerous globose ascospores, was also found at this site. This taxon was not found elsewhere in the DRBH, and is so unlike any other of which we are aware that it is described herein as a new genus and species (A. pamlicoensis). The number of rare and unusual species found at the one inland upland site that we inventoried highlights an important avenue for future study, and hints at the richness that lichen communities in upland hardwood forests once attained in the region.

Taxonomic Diversity

Taxonomic diversity (i.e., the number of species and infraspecific taxa) found at each site is summarized in Figure 5. Although taxonomic diversity values vary greatly across the DRBH (min. = 24, max. = 150), the average diversity was high (x̄ ± s = 67 ± 23) compared to other regions of the MACP (Lendemer and Allen 2014), with 57% (n = 24) of the sites hosting ≥80 taxa. Similarly, although collection effort was held constant across sites, the number of vouchers collected varied greatly (min. = 30, max. = 322), with an average of 100 ± 48 vouchers collected at each site. There is also a strong positive correlation between the number of collections and taxonomic diversity (Figure 6; R2 = 0.9149), which suggests that the team inventories conducted for this study did not result in repeated collection of the same species at the same sites, either by the same individual or between individuals. Of the 386 taxa encountered during the inventory, 97% (n = 360) were identifiable to species or infraspecific taxon, with the majority of the remaining 3% (n = 10) identified to genus and likely representing additional undescribed species.

Figure 5. 

Graph summarizing taxonomic diversity at each site across the Dare Regional Biodiversity Hotspot.


Figure 6. 

Scatter plot comparing collection effort (the number of collections made at each site) to taxonomic diversity on a per-site basis. Note that the outlier status of the highest diversity site may be due to the fact that it was the only site visited twice, once by the original inventory team, and then again by the participants of the Tuckerman Workshop.


Species Traits

Given the sampling strategy and large number of samples, we were curious to understand the frequencies of lichen traits in the study area. As such, we examined three commonly analyzed lichen traits (growth form, photobiont, and reproductive mode) as they relate to taxonomic diversity, the total number of specimens examined, and the total pooled occurrences for all species. Although limited, the examination of quantified traits performed for this study yielded a number of interesting results, which are summarized in Table 3.

Table 3. 

Summary of Dare Regional Biodiversity Hotspot taxonomic diversity and voucher/occurrence numbers, broken down by three commonly used lichen traits. Note that totals do not match voucher totals reported elsewhere herein, because they do not include unidentified specimens/taxa.


From the standpoint of growth form, the DRBH lichen biota is dominated by crustose lichens, which comprise fully 71% (272 taxa) of total lichen taxonomic diversity. Although intuitive, given the widely recognized diversity of crustose lichens, it is nonetheless surprising that the much more conspicuous and well-studied macrolichens comprise less than one-third (30%: 21% foliose lichens, 9% fruticose lichens) of the diversity. A similarly interesting result is that 68% (258 taxa) of the species in the DRBH reproduce sexually (i.e., produce apothecia or perithecia, and are presumed not to reproduce primarily through vegetative means), while 30% (113 taxa) reproduce through the dispersal of lichenized diaspores. The results for photobiont type are not unexpected for a temperate region with subtropical elements, as more than half (55%, 212 taxa) have a coccoid green algal photobiont and nearly a third (28%, 108 taxa) have Trentepohlia as a photobiont. The latter is a photobiont that is much more frequent in humid tropical regions compared to temperate or arctic regions (Nash et al. 1987, Matos et al. 2015). It is also noteworthy that small percentages of the species in the DRBH lack a photobiont (4%), are lichenicolous on other lichens (9%), or have a cyanobacterial photobiont (4%). Although the small number of cyanolichens present in the region may reflect a natural pattern, it is also possible that the number reflects declines as a result of habitat degradation and pollution (discussed in the “Overview: Vegetation and Lichen Diversity” section above). Finally, it is significant that the overall frequencies of different trait states in our diversity data were similar to those for the same trait states summarized from the numbers of vouchers and occurrences of each taxon. This suggests that our sampling was not biased toward any one trait state (i.e., the percentage of vouchers is negligibly different from the percentage of taxonomic diversity) and that the occurrence of a given trait state across the studied sites was negligibly different from the percentage of taxa with that trait state.

It should be noted that a logical extension of this aspect of our study would be to examine functional diversity, as well as trait distributions and correlates. We refrained from undertaking these analyses, because our dataset is taken from a relatively small geographic area. Instead, we will perform them on a much larger dataset covering the full MACP in a forthcoming study. It will be particularly interesting to see if the proportions of fruticose, asexual, cyanobacterial, or Trentepohlia-associated species observed in the DRBH are higher than in less intact and more degraded areas of the MACP. This is because previous studies of other regions have positively correlated numbers (diversity, frequency, and abundance) of taxa with those traits to higher-quality intact habitats (Marini et al. 2011, Stofer et al. 2006).

Comparison with Existing Data

It has already been highlighted above that one of the primary reasons for undertaking our inventory of the MACP, and by extension the DRBH, was that the region had previously been poorly studied. This is evidenced by the small number of floristic and taxonomic treatments that have covered the area, and by the fact that only 68 collections were documented from the DRBH in CNALH as part of the large-scale efforts to digitize lichen herbaria in the USA (see data deposited in Dryad). The majority of these were concentrated in highly visited areas of coastal barrier islands (Figure 1C). Although we acknowledge that there are almost certainly undigitized vouchers from the DRBH in other herbaria, there is no doubt that the collection bias and small number of vouchers reflect actual data gaps rather than data resource artifacts. With the substantial disparities between the pre- and poststudy data in mind, we nonetheless attempted to determine what impacts result from drawing conclusions from the prestudy data alone. This is salient, because large amounts of digitized lichen specimen data are now easily accessible online, and such data are actively being used without clear acknowledgement of their limitations.

First, we summarized and examined the CNALH raw data for DRBH lichens. Of the 68 vouchers that were available, 8 were undetermined and 24% (18 vouchers) were identified as 16 taxa that we did not encounter in our inventory. These are almost certainly misidentified, since available data indicate that the species do not occur within, or in some cases even near, the DRBH or the MACP (e.g., Lecanora albellula (Nyl.) Th. Fr. is so far unknown from eastern North America, Lepraria caesiella R.C. Harris does not occur south of the Delmarva Peninsula in the Coastal Plain, Parmotrema perlatum (Huds.) M. Choisy occurs much further inland, and all MACP specimens we have seen that are morphologically assignable to P. subtinctorium (Zahlbr.) Hale are actually P. neotropicum Kurok.). It is also likely that nomenclature updates of historical identifications resulted in several of these errors. For example: (a) Cladonia subcariosa Nyl. is broadly applied to a group including what we recognize as C. polycarpia G. Merr.; (b) Ramalina fastigiata (Pers.) Ach. would be updated to R. americana Hale, but that species is not known to occur in the DRBH; and (c) Bacidia atrogrisea (Delise) Körb. is a synonym of B. laurocerasi (Delise ex Duby) Zahlbr., but that species does not occur in the Coastal Plain. It is also possible that some of the vouchers are correctly identified. For instance, a voucher of Pseudocyphellaria crocata (L.) Vain could very well represent a collection of a species that is now extirpated from the DRBH. The above issues could be resolved by a loan of specimens from the relevant institution, and we assert that, before including CNALH data in lichen biodiversity assessments, any suspect specimens should be verified by physical examination. It should also be noted that, while we have opted not to examine these collections for this study, we did revise and examine all of the existing prestudy vouchers at NY, which were greater in number than those available in CNALH.

After summarizing the available data for the DRBH from CNALH, it was clear that sampling was strongly biased toward the coast, and the most diverse areas found in our study had not previously been sampled. Thus, any lichen threat assessment based on these data would have prioritized conservation efforts on the coastal barrier islands, which indeed are highly threatened, but ignored inland swamps that are similarly imperiled by sea-level rise, but host higher levels of diversity.

Lichen Community Similarity

Pairwise comparisons of the species compositions of the 49 sites inventoried for this study revealed a low degree of similarity between sites (Sørensen x̄ = 0.35 ± 0.12), suggesting a high degree of heterogeneity among sites. Indeed, the similarity values for sites in the DRBH are much lower than those obtained from a study on the nearby Delmarva Peninsula (Ray et al. 2015), a subregion of the MACP that hosts lower lichen diversity and has highly fragmented natural habitats (Lendemer and Allen 2014). A summary of taxonomic diversity and community similarity between the habitat types discussed in the “Overview: Vegetation and Lichen Diversity” section (above) is presented in Table 2. Overall, similarity between the lichen commuinites of different habitat types was low. Swamp forests and maritime forest uplands, on the other hand, were ∼60% similar. This relatively higher degree of similarity is likely to due to isolated swamp forests occurring within maritime forests on barrier islands (see “Distribution Patterns” below).

A high degree of lichen community heterogeneity is further supported by examination of the number of vouchers and number of occurrences per taxon, which are summarized in Figure 7. Remarkably, 23% (n = 88) of the taxa were collected only once, and fully 67% (n = 249) were collected 10 or fewer times (this includes multiple collection events at a single site). Thus, only 33% (n = 133) of the 386 taxa found in this study were collected more than 10 times, with 16% or 58 taxa which were collected between 11 and 20 times accounting for 77% of the taxa collected more than 10 times. The results of the occurrence data present a similar picture, with 27% (n = 100) taxa located at only 1 site, 74% (n = 275) taxa located at 10 or fewer sites, and only 29% (n = 107) taxa located are more than 10 sites. Again, regardless of whether one examines the total number of vouchers collected per taxon or the number of sites at which a given taxon occurred, more than half of the taxa were encountered only a small number of times, and a surprising number were collected or located only once.

Figure 7. 

Graphs summarizing the number of vouchers (left) and number occurrences (right) for species in the Dare Regional Biodiversity Hotspot.


Overlap between Collectors

Given that this was a study wherein a team of multiple experts inventoried sites of approximately equal size and with approximately equal effort, we examined a subset of the study sites to determine the amount of overlap between the species assemblages collected by different members of a team at a given site. Pairwise comparisons of the species assemblages collected by 3 team members (J.L.C., R.C.H., and W.R. Buck) at 24 sites revealed levels of similarity below 50% (Table 4). This supports anecdotal information and smaller studies (see Lendemer et al. 2013), suggesting that even within the small area reasonably surveyed for less than an hour, individuals will still collect very different assemblages of species. This is likely attributable to a combination of several factors, namely: (a) individually specific search patterns wherein one person may focus on the boles of trees, while another may focus on branches, roots, humus, or rocks; (b) individually specific taxonomic focuses (e.g., calicioid fungi, macrolichens, pyrenolichens, sterile lichens) that manifest even when conducting a complete biodiversity inventory; and (c) the actual existence of many more species in a small area than has generally been appreciated previously. Although the overlap between collectors may be greater in areas that host low levels of biodiversity, our results in particular highlight the importance of relying on trained experts to conduct biodiversity assessments and inventories. Results also highlight the value of a team approach, when the goal is to document total biodiversity. Indeed lichens are a nonmonophyletic group comprising species that belong to multiple fungal lineages. The expectation that a single individual will recognize and capture all lichen biodiversity is akin to expecting one mycologist to inventory all fungi or one biologist to document all invertebrate life.

Table 4. 

Overlap between species assemblages collected by three team members at 24 sites in the Dare Regional Biodiversity Hotspot expressed as the average ± SD of Sørensen similarity values obtained from all possible pairwise comparisons (n = 24 comparisons per collector pair).


Distribution Patterns

Examination of the distributions of 337 species within the DRBH and adjacent areas of the MACP (i.e., including the Chowan River Drainage and Great Dismal Swamp to the north, and Carteret County, Croatan National Forest, and the Pamlico Peninsula to the south) revealed distinct patterns and trends. The table summarizing species distributions, and the maps used in this part of this study, are both included in the data archive submitted to Dryad.

From the standpoint of overall distributions within the DRBH and adjacent areas of the MACP, assemblages of species were found to occur only in either inland swamp habitats or coastal maritime forests. These patterns reflect major differences in environmental conditions and vegetation (Schafale and Weakley 1990, Bellis 1995), and, although well documented in vascular plants (Kearney 1901, Bordeau and Oosting 1959, Griffith et al. 2002, Fleming 2012), they have been little studied in lichens. The phenomenon is illustrated by the genus Haematomma, wherein four species (H. accolens, H. americanum, H. flexuosum, and H. guyanense) occur only at inland sites, while one species is restricted to only maritime sites (H. persoonii). Of the taxa mapped for this study, 136 (41%) were found only at inland sites, 26 (8%) were found only in maritime sites, and 175 (52%) were found at both kinds of sites. Although 36 (11%) of the taxa were represented by only 1 occurrence on the distribution map, these singletons were disproportionately inland (29 taxa, 81% of singletons), while only a small number were maritime (4 taxa, 11% of singletons).

The large number of taxa found at both inland and maritime sites must also be interpreted in light of two factors. First, while some taxa are truly restricted to maritime habitats throughout their ranges in North America (Moore 1968, Harris 1995; Lendemer and Harris 2014a, 2015), many species are distributed only in such habitats at the northern edge of their range (e.g., Delmarva to South Carolina) and then also occur inland once they reach Florida (Brodo et al. 2008, Lendemer and Harris 2014b). Second, many of the taxa scored as present in both inland and maritime sites were mostly restricted to one region, with only one to several occurrences in the other region. Such outlier occurrences were of two types: (a) inland taxa present in mature deciduous maritime forests that occur on the inland sides of barrier islands (e.g., the majority of such singleton maritime occurrences of inland species are from an unusual hardwood forest at Kitty Hawk NERR); or (b) maritime taxa present in groups at a small number of unusual inland sites (e.g., the inland occurrences of Dirinaria confusa and Xyleborus nigricans). It is interesting to note that, although inland and maritime sites clearly host unique assemblages of taxa, these two groups of sites are less similar to each other (Sørensen x̄ ± s = 0.26 ± 0.09 between inland and maritime sites) when compared to the similarity among sites within each group (Sørensen x̄ ± s = 0.41 ± 0.11 for inland sites, 0.35 ± 0.11 for maritime sites).

In addition to the differences between inland and maritime habitats, an interesting pattern was noted, wherein some lichenicolous fungi were found only at inland sites, while their hosts were widely distributed throughout the study area (Figure 8). An example of this pattern is Gyalideopsis floridae, which is lichenicolous on members of the genus Parmotrema, particularly P. submarginale and P. subrigidum, both of which were widely distributed in the DRBH. In many cases, however, the distribution of the lichenicolous fungus does mirror that of the host, such as Buelliella trypethelii which was found only at inland sites where its host Bathelium carolinianum occurred, and Vouauxiella lichenicola, which occurs on different crustose lichens, but in the Coastal Plain is most common on Lecanora louisianae and found wherever that species occurs.

Figure 8. 

Comparison of lichenicolous fungus distributions between two hosts that occur throughout the Dare Regional Biodiversity Hotspot. A. Vouauxiella lichenicola (stars) occurs at both inland and maritime sites on Lecanora louisianae B. de Lesd. (white circles). B. Gyalideopsis floridae (stars) occurs only at inland sites, although the hosts, Parmotrema perforatum (white circles) and P. submarginale (gray circles), occur at both inland and maritime sites.


Floristic Elements

The Mid-Atlantic region of the Atlantic Coastal Plain is a biological transition zone that includes boreal or northern temperate and southern subtropical or tropical floristic elements in varying proportions, largely depending on latitude (Kearney 1901, Transeau 1903, Torrey 1937, Beaven and Oosting 1939, Ahti 1961, Dirig 1990, Lendemer and Knapp 2007). The DRBH fits well within this pattern, as evidenced by the taxa with continuous distributions in the Coastal Plain that have their northern limit (76 taxa, 21%) or southern limit (9 taxa, 3%) in the region. The large number of species with northern distributional limits in the DRBH supports the characterization of the region as hosting a biota dominated by southern or subtropical elements. Examples of taxa at the northern limits of their geographic ranges include members of genera that are particularly diverse in subtropical and tropical regions, such as Bactrospora (B. brevispora, B. carolinensis, B. lamprospora), Dirinaria (D. aegialita, D. confusa, D. picta), Fissurina (F. alligatorensis, F. columbina, F. incrustans, F. illiterata, F. scolecitis), Ocellularia s.l. (O. americana, O. praestans, O. sanfordiana), and Pyrenula (P. anomala, P. cruenta, P. microcarpa, P. microtheca, P. santensis). The northern distributional limit of foliicolous lichens in the Coastal Plain is also located in the DRBH. Asterothyrium decipiens occurs as sterile or pycnidiate thalli on the leaves of Persea Mill. in inland swamps, and Fellhanera bouteillei occurs as pycnidiate thalli on fronds of Sabal minor (Jacq.) Pers. in maritime forests. The populations of S. minor in the DRBH also represent the northern distributional limit of that species, and of the entire palm family (Arecaceae) in eastern North America (Tripp and Dexter 2006). It is interesting to note that both of the foliicolous lichens that are at the northern edge of their range in the DRBH only occur as sterile thalli with pycnidia.

Examples of taxa at the southern limits of their geographic ranges in the Coastal Plain include such northern temperate species as Anzia colpodes, Arthonia ruana, Flavoparmelia caperata, Lepraria harrisiana, Micarea peliocarpa, Phaeocalicium polyporaeum, and Ropalospora viridis. Many of the 230 taxa (66%) found in the DRBH that occur both north and south of the region are near the northern or southern limits of their geographic ranges within the Coastal Plain (Lendemer, unpubl. data). Since the only significant barriers for north–south lichen migration within the Coastal Plain are water bodies and availability of suitable habitat, the latter being a geologically recent anthropogenic constraint (Napton et al. 2010, Terando et al. 2014), understanding the present-day distributional limits of species establishes an important benchmark for future studies of global environmental change.

In addition to the temperate and subtropical elements present in the DRBH, there are two floristic elements that merit comment, one comprised of taxa endemic to the Coastal Plain of southeastern North America, and another comprised of tropical taxa that are disjunct from their ranges much further south in the Coastal Plain. The disjunct occurrence of subtropical and tropical species in the DRBH is not surprising, given that the region is characterized by a more southern lichen biota. Nonetheless, the majority of subtropical and tropical species found in the DRBH have been shown by our work elsewhere in the MACP to have continuous distributions in the Coastal Plain south of the DRBH (e.g., Lendemer and Harris 2015). Thus, taxa that are truly disjunct between the DRBH and the nearest populations often located >500 km to the south, such as Acrocordia gemmata, Bactrospora brevispora, Parmeliella pannosa, and Pyrgillus javanicus, are exceptions.

Endemic plants and animals have long been recognized as occurring both in the Coastal Plain and in its many subregions (e.g., Sorrie and Weakley 2001, Fleming 2012). In the MACP, the most iconic example may be the venus flytrap (Dionaea muscipula J. Ellis), narrowly endemic to a small portion of North Carolina and South Carolina (Sorrie and Weakley 2001). Lichens are no exception to this pattern of endemism, and while many species that occur in the Coastal Plain also occur in other tropical regions of the world, there are also many endemics and near endemics from a wide range of macrolichen and microlichen groups (Brodo 1968, Moore 1968, Harris 1995, Lücking et al. 2011, Lendemer and Harris 2015). Only two species discovered in our inventory appear to be endemic to the DRBH, A. pamlicoensis from Bull Neck Swamp and Lichenochora haematommatum parasitic on H persoonii from Hatteras Island. Otherwise, no other species are strictly endemic to the DRBH, although many are nearly endemic to the region. Examples of such taxa include Sticta deyana, which occurs in mature inland swamp forests, and Phaeographis oricola, which occurs in mature maritime forests on barrier islands.

Threats to the Lichen Biota

The remaining natural habitats in the MACP, and specifically in the DRBH, have been classified as threatened or endangered (e.g., Kirby-Smith and Barber 1979, Bellis 1995, Riggs and Ames 2003, Brown et al. 2005, Berman and Berquist 2007, Sallenger et al. 2012, Lorber and Rose 2015, Noss et al. 2015, US Army Corps of Engineers 2015). However, these assessments have not taken into account lichen diversity, an important and conspicuous component of the vegetation (see Lendemer and Allen 2014). The threats to DRBH lichen communities can be divided into two groups: those that have already had impacts in the past and will continue into the immediate future, and those that are projected to materialize in the future.

It is clear that the most significant impacts to the inland DRBH lichen communities have resulted from large-scale conversion of natural habitats for human uses (agriculture, sylviculture) as well as from resource extraction (e.g., Lorber and Rose 2015). Although the initial loss of suitable lichen habitat and substrates from these activities was immediate and substantial, long-term impacts of large-scale ditching and draining of water-logged swamps and peatlands have been far more pervasive and persistent, because hydrological regimes have been altered (Phipps et al. 1978, Kirby-Smith and Barber 1979, Daniel 1981, Ash et al. 1983). In the DRBH, like many areas of the MACP, natural habitats that remain primarily have poor soils unsuitable for agriculture (e.g., the sandhills), lie in difficult-to-access floodplains (e.g., bottomland swamps along rivers), or have persisted despite repeated attempts to alter them for human uses (e.g., swamps and peatlands). The scale and degree of changes associated with historical ditching and draining is difficult to appreciate in the present time. To place these effects in context, consider that attempts to drain the Great Dismal Swamp on the border of North Carolina and Virginia were initiated by George Washington in the late 1700s so that the 45,000-ha swamp that remains today is less than a third of its original area (see, e.g., Morse 1804, Kearney 1901, Simpson 1998). Although the importance of wetlands, swamps, and natural habitats generally has gained increasing recognition (Noe and Hupp 2005, 2009), the DRBH, like many other regions of the eastern USA continues to be affected by irreversible losses of habitat due to development and degradation of habitats by diverse forces (e.g., Figures 9A, 9C).

Figure 9. 

Examples of threats to the Dare Regional Biodiversity Hotspot lichen biota and habitats. A. Clear-cut logging of hardwood swamp parcel (cut 2014, inventory of adjacent protected area completed in 2013). B. Erosion of the shoreline. C. Development, construction, and maintenance of infrastructure, including highways and bridges. D. Sea-level rise, as exemplified by the conversion of healthy pocosins and swamp forests to marsh and, eventually, open water.


While the impacts of anthropogenic land use have been substantial and continue to persist as ongoing threats to biodiversity, the potential loss or irrevocable alteration of large areas through global climate change and sea level rise are major long-term issues for the DRBH (Riggs and Ames 2003, Sallenger et al. 2012). Much of the DRBH, including the lowest-lying swamp forests that host the highest lichen diversity in the MACP, is well within 1.5 m of current sea level, and is projected to be inundated by 2100 under the most conservative estimates (Figures 9B, 9D; Lendemer and Allen 2014). Although large areas of these unique habitats have been protected for the present, their continued existence into the future remains far from certain.


Although many of the natural communities in the DRBH cover large spatial areas, in some cases these are among the largest and best-preserved examples of those communities remaining (e.g., Lorber and Rose 2015). Furthermore, several of these communities are treated as critically imperiled, endangered, or rare within North Carolina and at a global scale (US Department of the Interior 2007, 2008). Thus, at both the natural community level and even landscape scale, the DRBH includes substantial protected areas that are vital to maintaining the integrity of the Atlantic Coastal Plain biome and the ecosystems services it provides.

In this context, the DRBH serves as the primary lichen biodiversity reservoir for the MACP (Lendemer and Allen 2014). Indeed, the region hosts the core ranges and largest populations of endemic, near-endemic, and regionally or globally rare or threatened species. As such, the DRBH lichen communities function as crucial, and in some cases the only, diaspore banks with which to establish new populations or attempt translocations. Research on the conservation and management of these source populations, as well as development of mitigation strategies to facilitate migration inland at pace with sea level rise, are an immediate concern and should be prioritized. This is particularly the case when one considers that, even under conservative estimates, sea level rise–related impacts to the DRBH are projected to be disproportionate compared to the rest of the MACP, and to occur within a relatively short time frame (Riggs and Ames 2003, Sallenger et al. 2012).

CONCLUSION That the lichens of both the MACP and the DRBH, two regions visited by millions of tourists annually and within several hours drive of major metropolitan areas, have received so little study previously, is remarkable. The data and results presented here not only provide the first comprehensive account of lichen biodiversity in the DRBH, but indeed for any large region of the Coastal Plain in southeastern North America, a biodiversity hotspot long known to host unique and diverse communities of plants and animals (James 1961, Estill and Cruzan 2001, Sorrie and Weakley 2001, Noss 2013, Noss et al. 2015). While a small number of accounts of southeastern Coastal Plain lichen biodiversity have been published, these have covered smaller areas with less-intensive sampling efforts (e.g., Hodkinson and Case 2008, Lücking et al. 2011) or been taxonomically incomplete (e.g., Moore 1968, Harris 1995).

The present study also summarizes diversity of an important component of obligate symbiont biodiversity in a biodiversity hotspot that has been delineated specifically based on the presence of high lichen diversity compared to the rest of the ecoregion. A critical avenue for further study involves determining whether the DRBH is also a hotspot for other groups of obligate symbiotic organisms.

It should be noted that many of the results presented here pertaining to patterns and trends of lichen biodiversity have previously been hypothesized, discussed anecdotally, or validated via studies that did not cover the entire lichen biota. Thus, this study constitutes an important quantitative analysis of lichen diversity and distributions in a biodiversity hotspot, and is, in-so-far as we are aware, unique for any area outside of Europe. As such, we hope that the methods, both sampling design and analyses, employed herein will serve as a useful model for future work in other areas of North America and abroad. In this manner, establishment of a robust body of scientific literature on lichen biodiversity, via studies the results of which can be directly compared, functions as a critical primary step to affect parity between the biodiversity data available for lichens and those of other macroscopic organisms, such as birds, mammals, and vascular plants.

ACKNOWLEDGMENTS This study and the authors were supported by NSF DEB-1145511 (to principal investigators J.C. Lendemer and R.C. Harris). We thank Erin Tripp for helpful discussion and encouragement in preparing the descriptive statistics portion of the manuscript. We thank Josef Hafellner and Toby Spribille for discussion of Albemarlea. We thank the following for their companionship in the field: J.L. Allen, W.R. Buck, T. McMullin, E. Tripp, and all the participants of the 2014 Tuckerman Workshop. We thank the staff of the Tranquil House Inn (Manteo, North Carolina) for their hospitality. The Coastal Studies Institute of the University of North Carolina (Wanchese, North Carolina) provided important logistical support. The staff and land managers of Alligator River National Wildlife Refuge, Bull Neck Swamp North Carolina State University Forest, Cape Hatteras National Seashore, Cape Lookout National Seashore, North Carolina National Estuarine Research Reserves, North Carolina Wildlife, Pine Island Audubon Sanctuary, Pocosin Lakes National Wildlife Refuge, and Tyrrell County Palmetto-Peartree Preserve provided access to protected areas and facilitating the issuring of permits to allow our work.



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APPENDIX I: New lichens, allied, and lichenicolous fungi encountered during the DRBH inventory

Albemarlea Lendemer & R.C. Harris gen. nov. Mycobank #815,461 Figures 10 and 11

Figure 10. 

Albemarlea parmlicoensis (all from the holotype). A. Gross morphology of the thallus illustrating apothecia and conspicuous macropycnidia. B. Detail of apothecium. C. Transverse section of apothecium in water. D. Intact ascus in KI. E. Dehiscing ascus in KI. F. Detail of ascospores in KI. Scale bars = 0.5 μm in A and B, 50 μm in C, and 20 μm in D–F.


Figure 11. 

Albemarlea pamlicoensis pycnidia (A–E, all from the holotype) and location of the type locality. A. Detail of macropycnidium with mass of conidia attached to the ostiole. B. Transverse section of macropycnidium in water. C. Macroconidia in phloxine. D. Transverse section of micropycnidium in water. E. Detail of micropycnidium also illustrating microconidia in water. F. Geographic distribution of A. pamlicoensis. Scale bars = 0.2 mm in A, 200 μm in B, 50 μm in D, and 20 μm in C and E.


Diagnosis. – A distinct genus of crustose lichenized ascomycetes with biatorine apothecia, Fuscidea-type asci that are polysporous and contain many hyaline, ellipsoid, simple ascospores, a coccoid photobiont, conspicuous superficial macropycnidia with narrowly fusiform two-celled hyaline macroconidia, and inconspicuous immersed micropycnidia with curved or bent rod-shaped simple hyaline microconidia.

TYPE: Albemarlea pamlicoensisLendemer & R.C. Harris

Etymology. – The epithet “Albemarlea” commemorates the Albemarle-Pamlico Peninsula of North Carolina, the only area of the Mid-Atlantic Coastal Plain (MACP) where the genus was found during our inventory. It concurrently commemorates the Albemarle Sound, which is a large body of water to the north of the Albemarle-Pamlico Peninsula. Bull Neck Swamp, where the only known population of this genus occurs, is a protected area that includes the largest undeveloped shoreline remaining on the Albemarle Sound.

Discussion. – The phenomenon of polyspory, or the production of more than eight ascospores per ascus, has been the subject of considerable interest in lichenology (Hafellner 1993, 1995; Reeb et al. 2004). Presumably this is due, in part, to the fact that researchers who study lichens are typically confronted with nearly uniform monotonous numbers of ascospores, so that when a species producing an unusual number is encountered, it immediately stands out as different. The occurrence of polyspory in lichen-forming fungi was recently summarized by Aptroot and Schumm (2012). Among the families and genera discussed by those authors, Maronea is similar to the new genus in having discoid apothecia (vs. perithecia or perithecioid-apothecia), Fuscidea-type asci, and numerous hyaline, simple ascospores per ascus (vs. smaller numbers greater than eight, such as 12, 24, 36, etc.). The new genus is readily distinguished from Maronea by its biatorine apothecia (vs. lecanorine apothecia with thalline margins) and the presence of conspicuous, superficial macropycnidia that produce hyaline, two-celled macroconidia, together with inconspicuous immersed micropycnidia that produce hyaline, curved, simple microconidia. In some respects, the apothecia and apothecial anatomy of the new genus resemble Piccolia, and, although untested with molecular data, we have tentatively considered that genus to be related to the Fuscidaceae based on ascus type (see Hafellner 1995). The apothecia of Piccolia contain orange, red, or yellow pigments that are strongly K+ purple or red, and the conidia (documented in P. conspersa (Fée) Hafellner and P. nannaria; Hafellner 1995, Lendemer and Harris 2014b) are simple and ellipsoid (vs. fusiform and septate or bent-fusiform and simple). As we are unaware of any other lichen that possesses the aforementioned characters, we here describe it as a new genus with a tentative placement in the Fuscideaceae pending further study with molecular methods.

Albemarlea pamlicoensis Lendemer & R.C. Harris sp. nov. Mycobank #815,462

Diagnosis. – A lichen-forming ascomycete with a crustose thallus, coccoid green photobiont, biatorine apothecia, Fuscidea-type asci containing many simple, ellipsoid, hyaline ascospores, conspicuous macropycnidia producing hyaline, narrowly fusiform, two-celled macroconidia, and inconspicuous micropycnidia immersed in the thallus and producing hyaline, bent or curved, rod-shaped simple macroconidia.

TYPE: USA, NORTH CAROLINA. Washington Co.: Bull Neck Swamp, south of Hufton Rd., 0.1–0.5 mi west of junction with Old North Bridge Rd., 35°57′50″N 76°26′33″W, 2 ft., upland mixed hardwood forest of Fagus, Quercus, Acer, Ilex with SymplocosVaccinium understory, 23 March 2013, on Fagus base, J.C. Lendemer et al. 36427 (NY!, holotype).

Description. – Thallus crustose, corticolous, greenish-gray, continuous, thin to thick, forming large continuous patches 7–15 cm in diameter, without soredia or isidia; prothallus indistinct, visible as a dark stain near the thallus margins. Apothecia biatorine, plane, circular in outline, sessile, reddish-brown in color, 0.3–0.5 mm in diameter, margins thin and slightly paler than the disc, quickly excluded with age; discs dark reddish-brown, epruinose; epihymenium hyaline to light tan, indistinct; hymenium 20–50 μm, hyaline, not inspersed; paraphyses slender, not or little branched, not distinctly expanded at the apices, hypothecium 100- to 230-μm thick, hyaline, inspersed with oil droplets; exciple 50- 70-μm thick, composed of thin radiating hyphae embedded in a thick gelatinous matrix, hyaline or except for a reddish-brown pigment in the outermost layer of cells. Asci short, clavate, Fuscidea-type, 50–80 × 20–40 μm; ascospores ellipsoid, simple, hyaline, many per ascus, thin walled, ca. 3.8–4.8 × 1.9–2.3 μm. Pycnidia of two types: macropycnidia abundant, conspicuous, raised above the thallus surface, walls reddish-brown, 250–300 × 50–70 μm; macroconidia forming a distinct white mass billowing out of the ostiole, narrowly fusiform, hyaline, one septate (rarely becoming two septate), 7.7–9.7 × 1.6–1.9 μm; micropycnidia sparse(?), inconspicuous, immersed in the thallus, walls hyaline, but becoming brown pigmented near the ostiole, 100–150 × 50–70 μm; microconidia rod-shaped, weakly to strongly bent and curved, simple, hyaline, 8–9.5 × 1.0–1.5 μm. Photobiont a coccoid green alga, cells 7–10 μm in diameter.

Chemistry. – No substances detected. Spot tests: K, C, KC, P, UV.

Etymology. – The epithet refers to the Pamlico Sound, a body of water located to the south of the type locality on the Albemarle-Pamlico Peninsula. The binomial A. pamlicoensis is intended to pay homage to the importance of the Albemarle-Pamlico Peninsula in serving as the primary reservoir for MACP lichen biodiversity.

Ecology and distribution. – Despite having surveyed more than 200 sites in the MACP, and revised thousands of Coastal Plain voucher specimens, the new species is known only from one robust population that was found growing on the base of a single mature American beech (Fagus grandifolia) at the type locality (Figure 11F). The type locality is an unusual inland, upland habitat with a relatively mature hardwood forest that is surrounded entirely by swamp forests that host very different vascular plant and lichen communities. We did not find other populations at any of the sites with similar natural communities (Merchants Millpond, North Carolina; Donnelly WMA, South Carolina).

Discussion. – In the field, A. parmlicoensis is most likely to be confused with sympatric crustose lichens that produce sporodochia (e.g., Dictyocatenulata alba, Xyleborus nigricans), because the macropycnidia that are raised well above the surface of the thallus tend to produce masses of macroconidia around the ostiole, thus giving the superficial appearance of sporodochia. When first located in the field, the new taxon was assumed to be the first fertile material of D. alba, a species easily distinguished by its Trentepohlia photobiont and sporodochia that are typically elevated on stalks (Lendemer and Harris 2004, Diederich et al. 2008).

Further study of the new taxon revealed the unusual combination of morphological characters, unlike any other polysporous lichen of which we are aware (see, e.g., Hafellner 1993, 1995). The Fuscidea-type asci suggest a relationship to Maronea; however, members of that genus have lecanorine apothecia and produce secondary compounds. As was suggested by colleagues (J. Hafellner and T. Spribille, pers. comm.), the apothecia of A. pamlicoensis are internally somewhat similar to Sarcosagium campestre (Fr.) Poetsch and Schied., although that species occurs on soil in northern temperate regions and noticeably differs from A. pamlicoensis in having asci with narrow I+ plugs in the tips (Hafellner 1995; Figure 10).

The internal anatomy of the apothecia, including the ascus type, of Albemarlea is also very similar to that of the genus Piccolia, particularly P. conspersa and P. nannaria. The new taxon is not likely to be confused with members of that genus on account of the very different conidia and the absence of orange or red, K+ red/purple pigments in the apothecia (Hafellner 1995, van der Broeck et al. 2013).

Arthonia agelastica R.C. Harris & Lendemer sp. nov. Mycobank #815,463 Figures 12 and 13

Figure 12. 

Arthonia agelastica (all from Harris 58367). A. Small group of ascomata associated with poorly developed apothecia of the host. B. Well-developed group of ascomata. C and D. Transverse section in water (C) and fuchsin (D) of apothecium (left) and pycnidium (right) on host thallus. E. Ascus with intact ascospores in iodine. F. Mature or postmature ascospores that have turned brown and are mounted in iodine. Scale bars = 0.5 mm in A, 0.25 mm in B, 100 μm in C and D, and 20 μm in E and F.


Figure 13. 

Geographic distribution of Arthonia agelastica.


Diagnosis. – A species of Arthonia Ach. s. lat. on Lecanora louisianae B. de Lesd. causing some bleaching or discoloration of the host thallus or less often causing no obvious damage. Ascomata light to dark brown, immersed, occurring in scattered to rarely ± confluent groups. Ascospores two (three) septate, macrocephalic, 13–14.7–16.7 × 5.2–6.0–7.5 μm, halonate.

TYPE: USA, NORTH CAROLINA. Tyrrell Co.: Pocosin Lakes National Wildlife Refuge, Frying Pan Boating Access, south of Frying Pan Rd., 6 mi east of junction with NC 94, 35°48′12″N 76°06′30″W, swamp forest of young mixed hardwoods (Acer, Liquidambar, Magnolia virginiana, Ilex) with sparse Taxodium, 10 December 2012, on Lecanora louisianae on fallen branch, R.C. Harris 58367 (NY!, holotype).

Description. – Ascomata immersed in thallus of Lecanora louisianae, in groups of few to many ascomata, light brown to dark brown, but sometimes discolored and blackish, epruinose, emarginate, 130- to 185-μm across, 100-μm high; epihymenium brown; hymenium colorless; hymenial gel I+ orangish, KI+ blue; paraphyses slender, weakly expanded at tips with brown caps; hypothecium colorless. Asci initially broadly clavate, more elongate at maturity, with tiny KI+ apical ring, with 8 spores; ascospores colorless becoming brown and warted in age, two (three) septate, macrocephalic, 13–14.7–16.7 × 5.2–6.0–7.5 μm, halonate. Pycnidia (seen only once), immersed, ±globose, with pale yellow brown wall, ca. 60-μm across; conidia bacillar, hyaline, 3.5–5 × 1.2–1.5 μm.

Etymology. – The epithet “agelastica” (=disposed to herd together) refers to the tendency of the ascomata to occur in discrete groups, or herds, on the thallus of the host.

Chemistry. – No substances detected. Spot tests: K, C, KC, P, UV.

Ecology and distribution. – Arthonia agelastica is evidently an obligate parasite on thalli of Lecanora louisianae, a crustose lichen that is common and widespread throughout the Coastal Plain of southeastern North America. The host typically occurs on the bark and branches of hardwood trees or shrubs, particularly in open swamp or coastal habitats.

It is notable that, although Lecanora louisianae is nearly ubiquitous from Delaware to Texas (and as far south as Hendry County in Florida), A. agelastica appears to have a considerably more restricted distribution (Figure 13). A survey of the 261 collections of L. louisianae held at NY located only five records in addition to those collected during our fieldwork on the Dare Peninsula. Of the 126 collections from Florida, only two were found to host A. agelastica. Similarly, only 3 collections of A. agelastica were found among the 64 collections of L. louisianae from North Carolina. Our search of the 71 additional collections of L. louisianae from other states in eastern North America failed to reveal any additional material of A. agelastica.

Discussion. – Arthonia agelastica is readily identifiable by its two-septate macrocephalic ascospores and ascomata that are immersed in the thallus of Lecanora louisianae. A number of other species of Arthonia have been reported to occur on various Lecanora species (Lawrey and Diederich 2011). Of these species, only A. subfuscicola (Linds.) Triebel has been reported from members of the L. subfusca group (in a broad sense), specifically from L. carpinea (L.) Vainio and L. pallida (Schreb.) Rabenh. (Triebel et al. 1991). A. subfuscicola differs from the new species in occurring mainly in the hypothecium of the apothecia of the host (vs. on the thallus), in having a dark hypothecium (vs. a hyaline one), and in having three-septate (vs. two-septate) ascospores (Triebel et al. 1991).

Additional specimens examined (all on thalli of Lecanora louisianae). – FLORIDA. Glades Co.: Ortona Cemetery, along SR 78, 1 mi west of CR 78A, 30 March 1998, on branch of Quercus, R.C. Harris 42119-A (NY). Levy Co.: Suwannee National Wildlife Refuge, Shell Mound County Park at west end of Co. Rd. 326, 3 December 1993, on branch, R.C. Harris 31477 (NY). NORTH CAROLINA. Carteret Co.: Cedar Island National Wildlife Refuge, south of Lola Rd. ∼1.3 mi southeast of junction with NC 12, 24 October 2012, on Ilex, J.C. Lendemer et al. 38416 (NY). Currituck Co.: Currituck Banks National Estuarine Research Reserve, west side adjacent to Currituck Sound, 0-1 mi north of terminus of NC 12 in Corolla, 14 April 2012, on Quercus, R.C. Harris 57272-A (NY); North River Game Land, west of Maple Rd., 0.5 mi north of intersection with US 158, 12 April 2012, on Acer, J.C. Lendemer 30715-A (NY). Dare Co.: Alligator River National Wildlife Refuge, west of Brier Hall Rd., 1.6 mi north of junction with US 64, 8 December 2012, on fallen branch, R.C. Harris 58113-A (NY); Alligator River National Wildlife Refuge, west of Buffalo City Rd., 1.2 mi south of US 64, 12 December 2012, on Acer, R.C. Harris 58623 (NY); Alligator River National Wildlife Refuge, southeast of junction of Spring Rd. and Navy Shell Rd., 19 March 2014, on Acer, J.C. Lendemer et al. 42658 (NY); Buxton Woods Coastal Reserve, southwest of terminus of Old Doctor Rd., west of Lookout Loop Trail, 18 August 2013, on Vitis, J.C. Lendemer 35850-A (NY); Cape Hatteras National Seashore, just west of Ramp 30, west of NC 12, north of Avalon, 19 March 2013, on Myrica, J.C. Lendemer 36309A (NY); Kill Devil Hills, Nags Head Woods Ecological Preserve, along Old Nags Head Woods Rd., 29 September 1993, on branches, W.R. Buck 24121-A (NY). Hyde Co.: Pocosin Lakes National Wildlife Refuge, south of New Lake Rd./SR1303, 7 mi northeast of junction with Higginsport Rd./SR 1302, 11 December 2012, on Acer, J.C. Lendemer 34871 (NY); Cape Hatteras National Seashore, north of NC 12, 0.25 mi west of Old Hammock Creek, 20 March 2014, on Ilex, J.C. Lendemer 42746 & E. Tripp (NY). Tyrrell Co.: Pocosin Lakes National Wildlife Refuge, Frying Pan Boating Access, south of Frying Pan Rd., 6 mi east of junction with NC 94, 10 December 2012, on twigs, W.R. Buck 60025 (NY), on upper branch of fallen Acer, R.C. Harris 58371-A (NY), on fallen branch, R.C. Harris 58393-A (NY). Washington Co.: Bull Neck Swamp, Deep Creek Rd., north of junction with Bear Lane, 23 March 2013, on Acer, E. Tripp et al. 4142 (NY); Bull Neck Swamp, south of Hufton Rd. 0.1–0.5 mi west of junction with Old North Bridge Rd., 23 Mar 2013, on Acer, J.C. Lendemer et al. 36471 (NY).

Arthonia hodgesii Lendemer & R.C. Harris sp. nov. Mycobank #815,464 Figures 14 and 15

Figure 14. 

Arthonia hodgesii (all from the holotype). A and B. Detail of fruiting bodies and infection occurring on thallus of Graphis lineola. C. Transverse section of ascoma in water. D. Transverse section of ascoma in K illustrating reaction of pigments. E and F. Ascus and ascospores in water. Scale bars = 500 μm in A and B, 50 μm in C and D, and 20 μm in E and F.


Figure 15. 

Geographic distribution of Arthonia hodgesii.


Diagnosis. – Differing from Arthonia graphidicola in having a brownish-orange epihymenium that is K+ magenta (vs. K+ dull olive) and an I+ blue (vs. I) hymenium.

TYPE: USA, GEORGIA. Doughtery Co.: Chickasawhatchee Wildlife Management Area, 31°29′25″N 84°25′07″W, pond cypress swamp forest, 10 November 2012, on Graphis lineola on branch of Morella cyrifera, M.F. Hodges 9228 (NY!, holotype).

Description. – Ascomata immersed in thallus of Graphis lineola, not producing any visible infection, light brownish-orange, slightly raised above the thallus surface, elongate and irregularly shaped 0.25–0.5 × 0.1–0.3 mm, often somewhat aggregated in a portion of the host thallus; epihymenium dark orange-brown pigmented, with the pigment turning K+ magenta but not dissolving; hymenium hyaline, I+ blue, KI+ greenish blue; hypothecium hyaline; paraphyses slender, weakly expanded at tips. Asci obovoid to obpyriform, without apical ring, 30–40 × 18–25 μm; ascospores three septate, hyaline, ±macrocephalic, becoming brownish and weakly roughened with age, I, (12.4–)12.8–13.6–14.3(−15) × (4.2–)4.5–5.1–5.8(−6.8) μm (n = 25). Pycnidia not seen.

Etymology. – The new species is named in honor of Malcolm Hodges, who, together with Sean Beeching, has contributed greatly to our knowledge of the lichen biota of Georgia. Both have an eye for unusual, small, or interesting species and are an asset to the field.

Chemistry. – Anthraquinone pigment in apothecia. Spot tests (pigmented portions): K+ magenta, KC, C, P, UV.

Ecology and distribution. – The new species is known only from two sites in the Coastal Plain of southeastern North America (Figure 15), where it was found growing in swamp forests on thalli of Graphis lineola on the bark of small hardwood trees and shrubs. Although the species is inconspicuous, it is readily visible and would likely have been detected if it were more widespread in the region where it occurs.

Discussion. – Arthonia hodgesii is most similar to A. graphidicola Coppins, a species described from Graphis scripta (L.) Ach. in Great Britain (Coppins 1989) and subsequently reported from France (Coste 1993) and Spain (Etayo and Diederich 1998). Although both species have similarly sized three-septate ascospores that are hyaline and macrocephallic, A. hodgesii differs from A. graphidicola in having an orange pigment in epihymenium that is K+ magenta (vs. a reddish-brown pigment that is K+ dull olive) and an I+ blue hymenium (vs. I) (fide Coppins 1989). The species also differ in their hosts, with A. graphidicola occurring on the primarily temperate species, G. scripta, and A. hodgesii occurring on G. lineola.

Arthonia diorygmae S. Joshi and Upreti is another lichenicolous species that was recently described from a host belonging to the family Graphidaceae in its traditional sense (Joshi et al. 2013). That species differs from A. hodgesii in numerous respects, including having circular ascomata and one-septate ascospores. The authors described A. diorygmae as having a brown to olive epihymenium that was “K+ slightly purplish” and, thus, it is unclear what type of pigment is present in the species. It should be noted that, in discussing their new species, as well as comparing it in a key to other lichenicolous species, Joshi et al. (2013) did not note the absence of K+ purple pigments in the epihymenium, as originally reported by Coppins (1989). Thus, those relying on the key published by Joshi et al. (2013) would likely incorrectly key A. hodgesii to A. graphidicola.

Recently, the circumscription of the Graphidaceae has been greatly expanded to include not only the entire Thelotremataceae, but also several other morphologically divergent families (Rivas Plata et al. 2012). Although we have elected to follow a more conservative taxonomy (Hodkinson 2012), it should be noted that Arthonia thelotrematis Coppins has been described from Thelotrema lepadinum (Ach.) Ach., a host that would now be placed in the same family as the hosts of A. graphidicola and A. hodgesii. As described by Coppins (1989), A. thelotrematis differs from A. hodgesii in having reddish-brown epihymenium that is K+ greenish, a similarly pigmented hypothecium (vs. hyaline in A. hodgesii and A. graphidicola), as well as slightly smaller ascospores (11–14 × 4.5–5.5 μm). Based on the available literature, A. hodgesii appears to be only the third species of Arthonia described from members of Graphidaceae in the traditional sense of that family.

Additional specimen examined. – USA, NORTH CAROLINA. Dare Co.: Alligator River National Wildlife Refuge, west of Whipping Creek Rd., 0.5 mi north of junction with Chip Rd., 23 March 2014, on Graphis lineola on dead sapling, R.C. Harris 60261-B (NY).

Arthonia stevensoniana R.C. Harris & Lendemer sp. nov. Mycobank #815,465 Figure 16

Figure 16. 

Arthonia stevensoniana morphology (A–E, all from the holotype), and geographic distribution. A. Gross morphology of infection in apothecia of Haematomma accolens. B. Detail of infection on apothecium of H. accolens. C. Transverse section of host apothecium with immersed ascoma, mounted in water. D. Ascus and ascospores in water. E. Transverse section of pycnidium in water. F. Geographic distribution of A. stevensoniana. Scale bars = 0.5 mm in A and B, 200 μm in C, and 20 μm in D and E.


Diagnosis. – Arthonia in hymenium of Haematomma accolens (Stirt.) Hillmann with brown hypothecium, dark green to green-black in KOH, hymenium I+ orange and ascospores 10–14 × 4–5 μm.

TYPE: USA, GEORGIA. Candler Co: Charles Harold TNC Preserve, 0–0.25 mi north of Salem Church Rd., west side of Stocking Head Creek, 32°25′01″N, 82°04′09″W, bottomland mixed hardwood forest (Nyssa, Acer, Quercus) with pine (Pinus), 22 December 2009, in hymenium of apothecia of Haematomma accolens on Acer, J.C. Lendemer et al. 21768 (NY!, holotype).

Description. – Ascomata in hymenium of apothecia of Haematomma accolens, black, flush with surface of host hymenium or slightly raised, 0.08- (immature) to 0.3-mm across, one to many per apothecium, sometimes coalescing so that entire disk of host is blackened; epihymenium brown; hymenium brown streaked, lower part dark greenish in KOH, I+ orange, KI+ greenish blue; hypothecium brown, dark greenish or greenish black in KOH; paraphyses mostly indistinct, thin, some weakly swollen at tip with a dark cap. Asci obovoid to obpyriform, without apical ring, 25–40 × 12–17 μm; ascospores one septate, hyaline, ±soleiform, becoming brownish and weakly roughened with age, upper cell slightly broader and longer than lower, with an I+ orangish sheath, (9.4–)10.1–11.1–12.1(−13.7) × (3.7–)3.9–4.3–4.8(−5.6) μm (n = 52). Pycnidia black, immersed in host hymenium, ca. 35–45 μm across, upper part of wall brown, lower part paler; conidia hyaline, narrowly fusiform, ca. 3.7–4.5(−5.2) × 1.1–1.6(−1.8) μm.

Etymology. – The epithet of the new species honors Robert Louis Stevenson (1850–1894), author of Treasure Island and Strange Case of Dr. Jekyll and Mr. Hyde. The reasoning for this eponymy is twofold; first the Arthonia is a pirate that takes over the apothecia of the host thallus, and second that the black spots on the red discs of Haematomma are reminiscent of the practice in Treasure Island of pirates giving the black spot as a threat of harm or death. It is also worth noting that Stevenson was a lover of islands, and traveled with his father throughout Scotland to examine the many lighthouses that his family had designed. Islands and lighthouses are iconic features of the region where the species grows, which was once home to many notorious pirates.

Ecology and distribution. – Arthonia stevensoniana appears to be endemic to the southeastern Coastal Plain of eastern North America, where it is known from a small number of inland swamp forest sites scattered across Georgia and North Carolina (Figure 16F). Interestingly, it is restricted to the apothecia of Haematomma accolens, a species that is common and widespread in portions of the Coastal Plain (Brodo et al. 2008). We suspect that the species is host specific and rare, given the frequency of Haematomma species in the region where it occurs, and the large amount of study that the genus has received (see below).

Discussion. – Species of the crustose lichen genus Haematomma, and the lichenicolous fungi that occur on them, have been the subject of considerable study and taxonomic treatments (see, e.g., Culberson 1963, Asahina 1964, Rogers 1982, Rogers and Bartlett 1986, Culberson et al. 1986, Kalb et al. 1995, Elix 2004, Nelsen et al. 2006, Brodo 2007, Brodo et al. 2008, Lumbsch et al. 2008). Similarly well studied are the lichenicolous species of Arthonia, which often form conspicuous infections on their host lichens and, thus, are routinely collected even by those not specializing in lichenicolous fungi (see, e.g., Santesson 1993, Grube et al. 1995, Hafellner 1995, Grube and Matzer 1997, Wedin and Hafellner 1998, Santesson et al. 2004). Given the robust body of literature devoted to these groups, and the large number of specimens of Haematomma that exist in herbaria, it was surprising to discover a previously unknown Arthonia that forms conspicuous black infections on the apothecia of Haematomma.

Among the lichenicolous fungi that occur on Haematomma, only Arthonia haematommatum Kalb and Hafellner from New Zealand also occurs in the hymenium of the host. However, that species differs from A. stevensoniana in having three-septate (vs. one-septate) ascospores, and in occurring on a different host with a different geographic distribution (H. accolens vs. H. alpinum R.W. Rogers and H. babingtonii A. Massal.; see Kalb et al. 1995). Although other lichenicolous fungi may blacken the apothecial discs of Haematomma species, none of these produce ascomata and have one-septate ascospores.

There are several species of Arthonia with one-septate ascospores that blacken the apothecia of other related lichen genera, such as Lecanora and Rhizoplaca. In addition to occurring on other hosts, two such species (A. apotheciorum (A. Massal.) Almq. occurring on members of the Lecanora dispersa group and A. Clemens (Tul.) Th. Fr. occurring on Rhizoplaca) differ from A. stevensoniana in having hyaline rather than brown hypothecia. The remaining species (A. galactinaria Leight. [=A. apotheciorum?] occurring on L. dispersa (Pers.) Sommerf., A. lecanorina (Almq.) R. Sant. Occurring on L. albella (Pers.) Ach.) have dark hypothecia and ascospores similar in size to those of A. stevensoniana, and thus must be separated based on their host preference. Although further study of this group is needed, it should be noted that neither L. albella nor L. dispersa are sympatric with Haematomma accolens in North America, or evidently with A. stevensoniana.

Additional specimens examined (all in the hymenium of apothecia of Haematomma accolens). USA, GEORGIA. Coffee Co.: Broxton Rocks Ecological Preserve, 9 mi northeast of Broxton, falls of Rocky Creek, 8 February 2003, on Quercus, R.C. Harris 47039 (NY). Pierce Co.: Little Satilla Wildlife Management Area, Knight Rd., ca. 1 mi east of Offerman between Zero Bay and Sixty Foot Branch of Little Satilla River, 21 December 2009, on Acer, J.C. Lendemer et al. 21533 (NY). USA, NORTH CAROLINA. Dare Co.: Alligator River National Wildlife Refuge, west of Whipping Creek Rd., 0.5 mi north of junction with Chip Rd., 23 March 2014, on dead sapling, R.C. Harris 60260 (NY). Tyrrell Co.: Palmetto-Peartree Preserve, northwest of junction of Loop Rd. and Canoe Pier Spur, 0.5 mi north of junction of Loop Rd. and Pot Licker Rd., 12 December 2012, on Acer, R.C. Harris 58567 (NY).

Lichenochora haematommatum R.C. Harris & Lendemer sp. nov. Mycobank #815,466 Figure 17

Figure 17. 

Lichenochora haematommatum morphology (A–E) and geographic distribution. A and B. Infection forming conspicuous galls on Haematomma persoonii (from Lendemer 36123). C. Transverse section of perithecium in IKI (from Buck 63107). D. Asci and ascospores in water (from Lendemer 36123). E. Hyaline ascospores (from Buck 63107). F. Brown ascospores (from Lendemer 36101). G. Geographic distribution of L. haematommatum. Scale bars = 0.5 mm in A and B, 200 μm in C, and 20 μm in D–F.


Diagnosis. – Lichenochora forming galls on the thallus and apothecia of Haematomma persoonii that contain few to many perithecia in various stages of maturity. Ascospores initially colorless, finally brown, broadly ellipsoid, one septate, constricted at the septum, (12–) 12.9–14.0–15.0(−16.4) × (7.1–)7.5–8.0–8.5(−9.3) μm, with a punctate perispore.

TYPE: USA, NORTH CAROLINA. Dare Co.: Cape Hatteras National Seashore, east of Lighthouse Visitor Center, 35°15′04″N, 75°31′29″W, elevation 7 ft., maritime forest of Pinus-Juniperus-Quercus with Ilex vomitoria understory, 18 March 2013, on H. persoonii on Quercus, J. C. Lendemer 36123 (NY!, holotype).

Description. – Lichenicolous fungus forming galls on thallus and apothecia of H. persoonii; galls concolorous with the host thallus, ±orbicular or irregular in shape, containing one to many perithecia, 0.2-mm across (single perithecium per gall) to 1.0-mm across (multiple perithecia per gall). Perithecia immersed to varying levels in galls, with only black tips visible, typically solitary but occasionally two to three perithecia becoming fused, black, pyriform, 125- to 130-μm wide, 150- to 200-μm tall, with black wall of elongated cells, 14- to 23-μm thick; periphyses present; hymenium filled with numerous oil droplets; paraphyses deliquescing. Asci ±cylindrical, uniformly thin walled; ascospores hayline, finally brown, occasionally weakly brown when still inside the ascus, broadly ellipsoid, one septate, ±constricted at the septum initially and becoming markedly so with age, (12–)12.9–14.0–15.0(−16.4) × (7.1–)7.5–8.0–8.5(−9.3) μm (n = 34), with a punctate perispore. Pycnidia not seen.

Etymology. – The epithet refers to the host genus Haematomma, as this appears to be the first member of Lichenochroa discovered on that host.

Ecology and distribution. – Lichenochora haematommatum is so far known only from two sites within a small area of Hatteras Island on the Outer Banks of North Carolina (Figure 17F). It is not uncommon at these sites, and is particularly abundant at the type locality where nearly every thallus of the host species appears to be infected. The restricted distribution of the new species does not appear to be an artifact of collection bias, as its host, H. persoonii, occurs throughout much of coastal southeastern North America and often dominates corticolous lichen communities on hardwood shrubs and branches in open, scrubby maritime forests. Despite having inventoried hundreds of sites in the southeastern Coastal Plain, and having made more than 100 collections of H. persoonii, the new species remains known only from Hatteras Island. Thus, it is possible that the species may be narrowly endemic to the area, or to the Carolinian Barrier Island ecoregion (see Griffith et al. 2002).

Discussion. – The genus Lichenochora (Phyllachoraceae) is easily recognized by its perithecia, hymenium that is obscured by oil droplets, deliquescing paraphyses, and uniformly thin-walled asci. Etayo and Navarro-Rosinés (2008) provided a key to the species of Lichenochora known at that time. In that key L. haematommatum comes closest to L. aipoliae Etayo, Nav.-Ros., and Coppins, a species described from Great Britain that has similar-sized, broad, ornamented, one-septate ascospores where the cells are of equal size, and which turn brown with age. The new species differs from L. aipoliae in having broader ascospores (7–9 μm vs. 5–7 μm) and in host preference (the crustose genus Haematomma vs. the foliose genus Physcia). Of the species described after 2008, L. physciicola (Ihlen and R. Sant.) Hafellner described from Sweden is most similar. Although the ascospores in that species are sized 11–13(−14) × 7–9 μm (fide Ihlen and Wedin 2005), they nonetheless differ from those of L. haematommatum in remaining hyaline (vs. turning brown) and in having a smooth perispore that is not ornamented (vs. a distinctly punctate ornamented perispore). Lichenochora physciicola further differs from L. haematommatum in its host, occurring on the foliose species Physcia dubia (Hoffm.) Lettau rather than crustose Haematomma.

Additional specimens examined (all on H. persoonii). USA, NORTH CAROLINA. Dare Co.: same locality as for the type, 18 March 2013, on Quercus, J.C. Lendemer 36092-A (NY), J.C. Lendemer 36101 (NY); Cape Hatteras National Seashore, trail from World War II Memorial, 0.5 mi west of Lighthouse Rd., 24 March 2014, on Quercus, W.R. Buck 63107 (NY), J.C. Lendemer 43159 (NY).

Megalaria alligatorensis Lendemer sp. nov. Mycobank #815,467 Figure 18

Figure 18. 

Megalaria alligatorensis morphology (A–E, from the holotype) and geographic distribution. A. Gross morphology of the thallus. B. Detail of apothecia. C–E. Transverse section in water (C), in water under polarized light (D), and after treatment in K (E). F. Geographic distribution of M. alligatorensis. Scale bars = 1.0 mm in A, 0.5 mm in B, and 50 μm C–E.


Diagnosis. – Similar to Megalaria albocincta and M. anaglyptica, but differing in its smaller and narrower ascospores.

TYPE: USA, NORTH CAROLINA. Hyde Co.: Alligator River National Wildlife Refuge, Chip Rd. 2 mi southwest of junction with Whipping Creek Rd., 35°38′40″N 75°58′42″W, 2 ft., pocosin dominated by Pinus and Gordonia, with sparse Acer and Magnolia virginiana, understory of Cyrilla, Ilex glabra, and Persea, 23 March 2014, on Cyrilla, J.C. Lendemer et al. 43124 (NY!, holotype).

Description. – Thallus crustose, corticolous, greenish-blue, forming small, circular patches 2–3 cm in diameter, areolate, without soredia or isidia; prothallus white, fibrous, poorly developed, and visible between the areoles, becoming immersed and darkened at the margin near the growing edge of the thallus; areoles small, ±dispersed to crowded and becoming confluent, initially globose and convex, but eventually becoming flattened. Apothecia biatorine, plane and flat, circular in outline and rarely becoming ±deformed, sessile, 0.4–1.0 mm in diameter; margins pale, waxy white, contrasting strongly with the coloration of the discs, becoming excluded with age; discs dark blue–black, epruinose; epihymenium 10- to 20-μm thick, blue-gray pigmented, K; hymenium 50- to 80-μm thick, hyaline, not inspersed; hypothecium 25- to 50-μm thick, upper portions pigmented purple, K+ distinctly blue, lower portions pigmented brownish, K+ more intense brown; exciple bilayered, comprised of an inner layer 60- to 100-μm thick, comprised of textura intricata, lightly brownish pigmented and K+ yellow, and an outer layer 40- to 60-μm thick, comprised of thick, gelatinized hyphae densely inspersed with POL+ crystals, without pigment and K. Asci Bacidia-type, cylindrical to clavate, eight-spored; ascospores narrowly ellipsoid, often weakly bent to one side, hyaline, one septate, thick walled, not halonate, 12–14 × 3.8–5.5 μm. Pycnidia not seen.

Chemistry. – Atranorin, zeorin, and fumarprotocetraric acid. Spot tests: K+ weak yellow, KC, C, P+ orange-red, UV.

Etymology. – The epithet refers both to the type locality in the Alligator River National Wildlife Refuge and, more generally, to the Alligator River region of North Carolina where the species occurs.

Ecology and distribution. – The new species is known only from the type locality (Figure 18F), where it was found growing on the bark of a hardwood tree (sweet bay [Magnolia virginiana L.]) and an ericaceous shrub (titi [Cyrilla racemiflora L.]) in dense shade in the understory of a pocosin. Due to the difficulty in accessing and traversing pocosin peatlands, it is possible that the species is more widespread in the region and has simply been overlooked previously.

Discussion. – Based on the production of zeorin and the presence of a distinctly bilayered exciple, Megalaria alligatorensis is most likely related to other tropical species that would have been classified in the genus Catillochroma Kalb by Kalb (2007). The circumscription of Megalaria Hafellner followed here is the pragmatic one adopted by Fryday and Lendemer (2010), wherein members of Catillochroma and Lopezaria Kalb and Hafellner were lumped within a single, broadly circumscribed genus pending further study with molecular data. To date, such studies have not been undertaken, and thus the new species is assigned to Megalaria in a broad sense. Among the species of Megalaria s.l., M. alligatorensis is distinctive on account of its chemistry (atranorin, zeorin, and fumarprotocetraric acid), relatively smooth esorediate thallus, apothecial pigmentation, including stark white margins, and small ascospores. It is most similar to M. albocincta (Degel.) Tønsberg, a species that was originally described from the Azores (Degelius 1941a) and subsequently reported from a single North American site in a high-elevation southern Appalachian spruce–fir forest (Degelius 1941b). Indeed, both M. albocincta and M. alligatorensis have similar apothecial pigmentation, esorediate thalli, and M. albocincta occasionally produces fumarprotocetraric acid as an accessory to atranorin and zeorin (while fumarprotocetraric acid was present in both specimens of M. alligatorensis examined) (Ekman and Tønsberg 1996). Nonetheless, M. alligatorensis can be distinguished from M. albocincta by its smaller ascospores (12–14 × 3.8–5.5 μm in M. alligatorensis vs. [13–]15–17 × 6.5–8.5 μm in M. albocincta fide Degelius [1941a]), the brown pigment in the hypothecium in M. alligatorensis, and by an apparent preference for hardwood substrates in M. alligatorensis rather than the coniferous substrates M. albocincta occurs on. It should be noted that Schumm and Aptroot (2013: 289) illustrated and described a sorediate specimen from Terceira in the Azores under the name M. albocincta. Although geographically proximal to the type locality of M. albocincta, the material differs from the published accounts of M. albocincta in having a sorediate thallus.

Megalaria anaglyptica (Kremp.) Fryday and Lendemer is a Brazilian esorediate species that also produces atranorin, zeorin, and fumarprotocetraric acid (Kalb 2007). That species is easily distinguished from M. alligatorensis by its thick, lumpy, granular thallus, an absence of purple pigment in the hypothecium, and by its larger ascospores (17–22 × 4–6 μm fide Kalb 2007). In treating M. anaglyptica, Kalb (2007) also mentioned the existence of material from Minas Gerais in Brazil that differed from the type in having smaller granules and smaller ascospores. Further study of that material should be undertaken in conjunction with study of M. alligatorensis.

Additional specimen examined. – Same locality as for the type, 19 March 2014, on Magnolia virginiana, J.C. Lendemer et al. 43134 (NY).

Minutoexcipula miniatoexcipula R.C. Harris & Lendemer sp. nov. Mycobank #815,468 Figures 19 and 20

Figure 19. 

Morphology and distribution of Minutoexcipula miniatoexcipula (all micrographs from the holotype). A–C. Detail of sporodochia on thallus (A and B) and ascomatal warts (C) of Pertusaria epixantha. D and E. Transverse sections of a sporodochium mounted in water. F. Geographic distribution of M. miniatoexcipula. Scale bars = 0.25 mm in A–C, 50 μm in D, and 20 μm in E.


Figure 20. 

A, conidiophores of Minutoexcipula miniatoexcipula. B. Conidia of M. miniatoexcipula. Scale bars = 20 μm. Both images from the holotype.


Diagnosis. – Differing from all known species of Minutoexcipula D. Hawksw. & V. Atienza in the dark orange–red–pigmented exciple and nonseptate conidia.

TYPE: USA, NORTH CAROLINA. Washington Co.: Bull Neck Swamp, Deep Creek Rd., north of junction with Bear Lane, 35°56′56″N 76°24′02″W, 1 ft., swamp forest with Chamaecyparis, Taxodium, and mixed hardwoods (Acer, Magnolia virginiana, Persea) with Lyonia-Ilex glabra understory, 23 March 2013, on Pertusaria epixantha on large Magnolia virginiana, J.C. Lendemer et al. 36395 (NY!, holotype).

Description. – Conidiomata sporodochia-like, on thallus and warts of Pertusaria epixantha, not usually causing any evident damage, but occasionally occuring on hosts where the thallus has become degraded, presumably by the infection, black, discoid, slightly constricted at base, 50- to 150-μm across, ca. 50-μm tall; exciple pigmented orange red, ±unchanged in KOH, ca. 10-μm thick, composed of relatively few ±rounded cells; conidiophores 7- to 14-μm long; conidia dark brown, obpyriform with one end truncate, 4.7–5.3–5.8 × 3.0–3.3–4.0 μm.

Etymology. – The species is named for the distinctive orange red color of the exciple, “miniatus” plus “excipulum.”

Ecology and distribution. – Minutoexcipula miniatoexcipula is currently known from two disjunct clusters of populations in inland swamp forests of the MACP in North and South Carolina (Figure 19F). This disjunct distribution does not appear to be an artifact of collection bias, as our inventory did not detect the species in the intervening area. The presence of the species in these two areas may reflect the availability of large areas of relatively high-quality intact habitat found there. To date, all of the known populations have been found on the thallus and warts of Pertusaria epixantha, which is widespread and common in southeastern North America, including in the area between the two known clusters of populations of M. miniatoexcipula. In several cases, it was not possible to determine the host lichen with certainty, because the thallus was small, sterile, or highly degraded by the infection of the Minutoexcipula. Nonetheless, in all such cases, the host appeared to be P. epixantha and not another species of Pertusaria.

Discussion. – The red color of the exciple and the nonseptate conidia found in the new species have not been reported for the genus Minutoexcipula (Atienza and Hawksworth 1994, Diederich 2003). Nonetheless, the elongate, septate conidiophores, presence of an exciple, and occurrence on Pertusaria indicate that it should be included in Minutoexcipula. While we cannot discount the possibility that the red coloration of the exciple is a result of an interaction with the host, the species is otherwise easily distinguished by the small, nonseptate conidia. With the description of the new species here, there are three species of Minutoexcipula that occur on members of the genus Pertusaria. However, in our experience, they are confined to separate, albeit morphologically similar, host species in eastern North America. While M. miniatoexcipula is a parasite of P. epixantha, M. mariana is parasitic on P. pustulata and M. tuckerae is parasitic on P. texana.

Additional specimens examined. USA, NORTH CAROLINA. Dare Co.: Alligator River National Wildlife Refuge, west of Buffalo City Rd., 1.2 mi south of US 64, 12 December 2012, on P. epixantha on Acer, W.R. Buck 60160 (NY); Alligator River National Wildlife Refuge, west of Whipping Creek Rd. 0.5 mi north of junction with Chip Rd., 23 March 2014, on P. epixantha on dead Lyonia branch, W.R. Buck 63061 (NY), on P. epixantha on Ilex, R.C. Harris 60264 (NY). Tyrrell Co.: Alligator River Game Land, Middle Rd. 0–0.25 mi northeast of US 64, 1.8 mi northwest of Alligator, 22 March 2014, on P. epixantha on Acer, W.R. Buck 63049 (NY), R.C. Harris 60240A (NY). USA, SOUTH CAROLINA. Berkeley Co.: Francis Marion National Forest, vicinity of Pitch Landing at terminus of FS 192, 6 December 2013, on P. epixantha on fallen branch, W.R. Buck 62099 (NY); Francis Marion National Forest, FS 204F, 0.25 mi south of McConnel's Landing, 3 December 2013, on Pertusaria sp. on Quercus, J.C. Lendemer et al. 40946 (NY). Charleston Co.: Francis Marion National Forest, Buck Hall Recreation Area, Palmetto Trailhead at terminus of FS 242, 1 December 2013, on P. epixantha on Quercus, W.R. Buck 61724 (NY); Francis Marion National Forest, Wambaw Swamp, east of Elden Rd./ FS C-10-217, 0.3 mi south of junction with Victor Lincoln Rd./ FS C-10-154, 1 December 2013, on P. epixantha on Acer, W.R. Buck 61807 (NY); Francis Marion National Forest, Wambaw Swamp Wilderness, Wambaw Swamp, at bridge on Elden Rd./FS C-10-217, 0.4 mi north of junction with FS 217A, 1 December 2013, on Pertusaria sp. on Acer, J.C. Lendemer et al. 40270 (NY). Colleton Co.: Donnelley Wildlife Management Area, 0.2 mi southwest of Main Rd., 0.7 mi north of junction with Blocker Run Rd., 18 December 2013, on P. epixantha on Nyssa, J.C. Lendemer et al. 41710 (NY).

Trichosphaerella buckii R.C. Harris & Lendemer sp. nov. MycoBank #815,469 Figure 21

Figure 21. 

Trichosphaerella buckii (all from the holotype). A. Detail of infection with emergent perithecia. B. Transverse section of perithecium in water. C. Mount of perithecium in water viewed from above. D. Detail of ostiolar setae in water. E. Asci and intact ascospores. F. Part-spores. Scale bars = 0.5 mm in A, 100 μm in B and C, and 20 μm in D–F.


Diagnosis. – Trichosphaerella E. Bommer, M. Rousseau, and Sacc. with perithecia immersed in the thallus of Punctelia rudecta. Perithecia conical, brown, with unbranched septae around the ostiole. Ascospores breaking into 16 part spores. Part spores colorless, tetrahedral, with papillae at the apices.

TYPE: USA, NORTH CAROLINA. Tyrrell Co.: Alligator River Game Land, Middle Rd. 0.0.25 mi northeast of US 64, 1.8 mi northwest of Alligator, 35°55′34″N, 76°08′05″W, 0 m, mixed hardwood (Nyssa, Acer, Magnolia virginiana)–Taxodium swamp forest, on Punctelia rudecta on trunk of Acer, 22 March 2014, W.R. Buck 63057 (NY!, holotype).

Description. – Perithecia parasitic or saprophytic(?), immersed in moribund thallus of Punctelia rudecta, conical, ca. 0.1- to 0.2-mm across, 0.8- to 0.17-mm tall, with erumpent apex; perithecial wall prosoplectenchymatous, entire, olivaceous, greenish around the ostiole, ca. 15-μm thick; erumpent ostiolar region with brown, unbranched setae, ca. 15–25 × 4–4.5 μm long; ostiolar region periphysate; paraphyses sparse, thin walled, unbranched, ca. 2.5- to 3.0-μm wide; hymenial gel I. Asci cylindrical, thin walled, without tholus, I, 50–65 × 11–13 μm, with 16 part spores; part spores hyaline, tetrahedral with papillate apices, 4- to 6-μm across (including papillae). Pycnidia not seen.

Etymology. – The epithet honors William R. Buck (b. 1950), astute collector of minute lichens and lichenicolous fungi who contributed greatly to our inventory of the DRBH and the MACP. Many species have been elevated from obscurity via formal scientific description as a result of collections he has made.

Ecology and distribution. – Trichosphaerella buckii is known only from the type collection, found growing on Punctelia rudecta in the Coastal Plain of eastern North Carolina (Figure 22). Considering that the host is common and widespread, both in the Coastal Plain and, more generally, throughout temperate eastern North America, it is possible that new species may prove to be more widespread.

Figure 22. 

Geographic distribution of Trichosphaerella buckii.


Discussion. – Trichosphaerella buckii is easily recognized by the immersed, erumpent, setose perithecia, and the asci with 16 apically papillate tetrahedral part spores. This is the first report of a lichenicolous species of Trichosphaerella. The genus is a member of the Niessliaceae, distinguished from Niesslia Auersw. by the eight initial ascospores separating into 16 part spores (Samuels and Barr 1997, Rossman et al. 1999). Although other species placed in the genus are saprophytic, T. goniospora Döbbeler et al. (2015) with similar tetrahedral part spores was recently described from a liverwort. Niesslia tetrahedrospora Etayo, which was described from the lichen Dichosporidium nigrocinctum (Ehrenb.) G. Thor by Etayo (2002) has similar part spores and could also be placed in Trichosphaerella. Both N. tetrahedrospora and T. goniospora differ from the new taxon in host, in having superficial perithecia, and in lacking papillae on the spore apices.

APPENDIX II: Checklist of the Lichens and Allied Fungi of the Dare Regional Biodiversity Hotspot (DRBH)

The checklist presented below is arranged alphabetically by genus and species. Taxa not identified to species are largely excluded from the list, as are unpublished names of new species for which descriptions are currently in preparation. When partially identified taxa, or unpublished names, are included in the list, it is because the taxon is common enough to be frequently encountered in the study area. Throughout the list, lichenicolous fungi are denoted by an asterisk (*) before the name, and nonlichenized fungi that are treated with lichens are denoted with a plus symbol (+) before the name. Taxonomy and taxonomic authorities largely follow Esslinger (2014), and deviations from that work reflect the opinions of the authors.

*Abrothallus parmotrematis Diederich

Acanthothecis leucoxanthoides Lendemer

Acanthothecis mosquitensis (Tuck.) E. A. Tripp & Lendemer

Acanthothecis paucispora Lendemer & R.C. Harris

Acrocordia gemmata (Ach.) A. Massal.

Agyrium rufum (Pers.) Fr.

Amandinea langloisii Marbach

Amandinea milliaria (Tuck.) P. May & Sheard

Amandinea polyspora (Willey) E. Lay & P. May

Amandinea punctata (Hoffm.) Coppins & Scheid.

Anaptychia palmulata (Michx.) Vain.

Anisomeridium anisolobum (Müll. Arg.) Aptroot

Anisomeridium biforme (Borrer) R.C. Harris

Anisomeridium biformoides R.C. Harris

Anisomeridium polypori (Ellis & Everh.) M. E. Barr

Anthracothecium nanum (Zahlbr.) R.C. Harris

Anzia colpodes (Ach.) Stizenb.

Anzia ornata (Zahlbr.) Asahina

*Arthonia agelastica R.C. Harris & Lendemer

Arthonia albovirescens Nyl.

Arthonia anglica Coppins

Arthonia cinnabarina (DC.) Wallr.

Arthonia hodgesii Lendemer & R.C. Harris

Arthonia interveniens Nyl.

+Arthonia quintaria Nyl.

Arthonia ruana A. Massal.

Arthonia rubella (Fée) Nyl.

*Arthonia stevensoniana R.C. Harris & Lendemer

Arthonia susa R.C. Harris & Lendemer

+Arthopyrenia cinchonae (Ach.) Müll. Arg.

+Arthopyrenia taxodii R.C. Harris

Asterothyrium decipiens (Rehm) R. Sant.

Bacidia diffracta S. Ekman

Bacidia helicospora S. Ekman

Bacidia heterochroa (Müll. Arg.) Zahlbr.

Bacidia schweinitzii (Fr. ex Tuck.) A. Schneid.

Bacidina sp. This is a common species in the DRBH and MACP that occurs on the bark of hardwoods and can be recognized by its pale-brownish apothecia that lack internal pigments, lack POL+ crystals, and the absence of secondary chemistry.

Bacidina crystallifera S. Ekman

Bacidina egenula (Nyl.) Vězda

Bacidina varia S. Ekman

Bactrospora brevispora R.C. Harris

Bactrospora carolinensis (Ellis & Everh.) R.C. Harris

Bactrospora lamprospora (Nyl.) Lendemer

Bathelium carolinianum (Tuck.) R.C. Harris

Brigantiaea leucoxantha (Spreng.) R. Sant. & Hafellner

Buellia curtisii (Tuck.) Imshaug

Buellia elizae (Tuck.) Tuck.

Buellia imshaugiana R.C. Harris

Buellia stillingiana J. Steiner

Buellia vernicoma (Tuck.) Tuck.

Buellia wheeleri R.C. Harris

*Buelliella minimula (Tuck.) Fink

*Buelliella trypethelii (Tuck.) Fink

Bulbothrix isidiza (Nyl.) Hale

Bulbothrix scortella (Nyl.) Hale

Byssoloma leucoblepharum (Nyl.) Vain.

Byssoloma meadii (Tuck.) S. Ekman

Caloplaca camptidia (Tuck.) Zahlbr.

Caloplaca feracissima H. Magn.

Caloplaca flavorubescens (With.) J.R. Laundon

Candelaria concolor (Dicks.) Stein

Candelariella xanthostigmoides (Müll. Arg.) R.W. Rogers

Canoparmelia amazonica (Nyl.) Elix & Hale

Canoparmelia caroliniana (Nyl.) Elix & Hale

Catinaria atropurpurea (Schaer.) Vězda & Poelt

Chaenotheca hygrophila Tibell

Chaenothecopsis debilis (Sm.) Tibell

Chaenothecopsis nana Tibell

Chaenothecopsis pusilla (Ach.) A.F.W. Schmidt

Chaenothecopsis pusiola (Ach.) Vain.

Chrysothrix chamaecyparicola Lendemer

Chrysothrix xanthina (Vain.) Kalb

Cladonia atlantica A. Evans

Cladonia beaumontii (Tuck.) Vain.

Cladonia caespiticia (Pers.) Flörke

Cladonia didyma var. vulcanica (Zoll. & Moritzi) Vain.

Cladonia evansii Abbayes

Cladonia incrassata Flörke

Cladonia leporina Fr.

Cladonia macilenta var. bacillaris (Genth) Schaer.

Cladonia ochrochlora Flörke

Cladonia parasitica (Hoffm.) Hoffm.

Cladonia peziziformis (With.) J.R. Laundon

Cladonia polycarpia G. Merr.

Cladonia ramulosa (With.) J.R. Laundon

Cladonia rappii A. Evans

Cladonia santensis Tuck.

Cladonia subradiata (Vain.) Sandst.

Cladonia subtenuis (Abbayes) Mattick

Coccocarpia erythroxyli (Spreng.) Swinsc. & Krog

Coccocarpia palmicola (Spreng.) Arv. & D.J. Galloway

Coenogonium luteum (Dicks.) Kalb & Lücking. Note. – This name is used here in a broad sense for a taxon that is common and widespread in the southeastern Coastal Plain, but that is strongly disjunct from the typical range of the species in northern temperate or boreal regions (see Brodo et al. 2001). Specimens referred to this species in the MACP likely belong to a separate species; however, its distinction from C. luteum requires extensive additional study beyond the scope of the present work.

Collema pulcellum Ach. var. leucopeplum (Tuck.) Degel.

Collema subflaccidum Degel.

*Coniambigua phaeographidis Etayo & Diederich

Coniarthonia pyrrhula (Nyl.) Grube

*Cornutispora triangularis Diederich & Etayo

Crespoa crozalsiana (Harm.) Lendemer & B.P. Hodk.

Cresponea flava (Vain.) Egea & Torrente

*Dactylospora inquilina (Tuck.) Hafellner

Dendriscocaulon intricatulum (Nyl.) Henssen

*Didymocyrtis melanelixiae (Brackel) Diederich, R.C. Harris & Etayo

Dirinaria aegialita (Ach.) B.J. Moore

Dirinaria confusa D.D. Awasthi

Dirinaria picta (Sw.) Schaer. ex Clem.

Dyplolabia afzelii (Ach.) A. Massal.

Enterographa anguinella (Nyl.) Redinger

Epigloea pleiospora Döbbeler

*Etayoa trypethelii (Flakus & Kukwa) Diederich & Ertz

Fellhanera bouteillei (Desm.) Vězda

Fissurina alligatorensis Lendemer & R.C. Harris

Fissurina columbina (Tuck.) Staiger

Fissurina cypressi (Müll. Arg.) Lendemer

Fissurina illiterata (R.C. Harris) Lendemer

Fissurina incrustans Fée

Fissurina insidiosa C. Knight & Mitt.

Fissurina scolecitis (Tuck.) Lendemer

Fissurina subnitidula (Tuck.) Staiger

Flavoparmelia caperata (L.) Hale

Fuscopannaria leucosticta (Tuck.) P.M. Jørg.

Gassicurtia acidobaeomyceta Marbach

Glyphis cicatricosa Ach.

Glyphis scyphulifera (Ach.) Staiger

Graphis cincta (Pers.) Aptroot

Graphis crebra Vain.

Graphis desquamescens Fée

Graphis duplicata Ach.

Graphis endoxantha Nyl.

Graphis furcata Fée

Graphis handelii Zahlbr.

Graphis intermedians Vain.

Graphis inversa R.C. Harris

Graphis lineola Ach.

Graphis pinicola Zahlbr.

Graphis scripta (L.) Ach.

Graphis striatula (Ach.) Spreng.

Graphis tenella Ach.

Graphis vittata Müll. Arg.

Gyalideopsis buckii Lücking, Sérus., & Vězda

*Gyalideopsis floridae Etayo & Diederich

Haematomma accolens (Stirt.) Hillmann

Haematomma americanum Staiger & Kalb

Haematomma flexuosum Hillmann

Haematomma guyanense Staiger & Kalb

Haematomma persoonii (Fée) A. Massal.

Hafellia sp. This is a species that is infrequent, but widespread, in coastal maritime forests throughout the DRBH. It does not appear to be one of the species yet known from North America; however, further study is needed before it can be described.

Heterodermia albicans (Pers.) Swinsc. & Krog

Heterodermia casarettiana (A. Massal.) Trevis.

Heterodermia crocea R.C. Harris

Heterodermia leucomelos (L.) Poelt

Heterodermia obscurata (Nyl.) Trevis.

Hypotrachyna cryptochlora (Vain.) D. Hawksw. & A. Crespo

Hypotrachyna horrescens (Taylor) Swinsc. & Krog

Hypotrachyna livida (Taylor) Hale

Hypotrachyna minarum (Vain.) Krog & Swinsc.

Hypotrachyna osseoalba (Vain.) Y.S. Park & Hale

Hypotrachyna spumosa (Asahina) Krog & Swinsc.

*Intralichen lichenum (Diederich) D. Hawksw. & M.S. Cole

Lecanora caesiorubella Ach. subsp. glaucomodes (Nyl.) Imshaug & Brodo

Lecanora chlarotera Nyl.

Lecanora cinereofusca H. Magn.

Lecanora cupressi Tuck.

Lecanora floridula Lumbsch

Lecanora hybocarpa (Tuck.) Brodo

Lecanora imshaugii Brodo

Lecanora louisianae B. de Lesd.

Lecanora nothocaesiella R.C. Harris & Lendemer

Lecanora strobilina (Spreng.) Kieff.

Lecanora subpallens Zahlbr.

Leiorreuma explicans (Fink) Lendemer

Leiorreuma sericeum (Eschw.) Staiger

Lepraria aurescens Orange & Wolesley

Lepraria finkii (B. de Lesd.) R.C. Harris

Lepraria friabilis Lendemer, K. Knudsen, & Elix

Lepraria harrisiana Lendemer

Lepraria hodkinsoniana Lendemer

Lepraria vouauxii (Hue) R.C. Harris

Leptogium austroamericanum (Malme) C.W. Dodge

Leptogium azureum (Sw.) Mont.

Leptogium corticola (Taylor) Tuck.

Leptogium cyanescens (Rabenh.) Körb.

*Lichenochora haematommatum R.C. Harris & Lendemer

*Lichenoconium cargillianum (Linds.) D. Hawksw.

*Lichenoconium lecanorae (Vouaux) Dyko & D. Hawksw.

*Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw.

Lobaria ravenelii (Tuck.) Yoshim.

Loxospora confusa Lendemer

Maronea polyphaea H. Magn.

Mazosia carnea (Eckfeldt) Aptroot & M. Cáceres

Megalospora pachycheila (Tuck.) Sipman

Megalospora porphyritis (Tuck.) R.C. Harris

*Melanographa tribulodes (Tuck.) Müll. Arg.

*Merismatium sp. This taxon was found growing on a thallus of Rinodina maculans in an inland swamp. As only a single small specimen was available, we have refrained from studying it further until additional material is located.

Micarea chlorosticta (Tuck.) R.C. Harris

Micarea micrococca (Körb.) Gams ex Coppins

Micarea neostipitata Coppins & P. May

Micarea peliocarpa (Anzi) Coppins & R. Sant.

Micarea prasina Fr.

*Minutoexcipula mariana V. Atienza

*Minutoexcipula miniatoexcipula R.C. Harris & Lendemer

*Minutoexcipula tuckerae V. Atienza & D. Hawksw.

*Muellerella lichenicola (Sommerf.) D. Hawksw.

Multiclavula mucida (Fr.) R.H. Petersen

Mycocalicium subtile (Pers.) Szatala

+Mycoporum eschweileri (Müll. Arg.) R.C. Harris

+Mycoporum lacteum (Ach.) R.C. Harris

Myelochroa aurulenta (Tuck.) Elix & Hale

Nadvornikia sorediata R.C. Harris

*Nectriopsis rubefaciens (Ellis & Everh.) M.S. Cole & D. Hawksw.

Ocellularia americana Hale

Ocellularia praestans (Müll. Arg.) Hale

Ocellularia sanfordiana (Zahlbr.) Hale

Ochrolechia africana Vain.

*Opegrapha anomea Nyl.

Opegrapha corticola Coppins & P. James

Opegrapha varia Pers.

Opegrapha viridis (Ach.) Behlen & Desberger

Opegrapha vulgata Ach.

Pannaria lurida (Mont.) Nyl. subsp. quercicola P.M. Jørg.

Pannaria tavaresii P.M. Jørg.

Parmeliella pannosa (Sw.) Müll. Arg.

Parmeliopsis subambigua Gyeln.

Parmotrema dilatatum (Vain.) Hale

Parmotrema gardneri (C.W. Dodge) Sérus.

Parmotrema hypoleucinum (J. Steiner) Hale

Parmotrema hypotropum (Nyl.) Hale

Parmotrema internexum (Nyl.) Hale

Parmotrema madagascariaceum (Hue) Hale

Parmotrema mellissii (C.W. Dodge) Hale

Parmotrema neotropicum Kurok.

Parmotrema perforatum (Jacq.) A. Massal.

Parmotrema praesorediosum (Nyl.) Hale

Parmotrema rampoddense (Nyl.) Hale

Parmotrema reticulatum (Taylor) M. Choisy

Parmotrema subisidiosum (Müll. Arg.) Hale

Parmotrema submarginale (Michx.) DePriest & B. Hale

Parmotrema subrigidum Egan

Parmotrema tinctorum (Nyl.) Hale

Parmotrema ultralucens (Krog) Hale

Parmotrema xanthinum (Müll. Arg.) Hale

Peltigera neopolydactyla (Gyeln.) Gyeln.

Pertusaria epixantha R.C. Harris

Pertusaria neoscotica I.M. Lamb

Pertusaria obruta R.C. Harris

Pertusaria paratuberculifera Dibben

Pertusaria propinqua Müll. Arg.

Pertusaria pustulata (Ach.) Duby

Pertusaria sinusmexicani Dibben

Pertusaria subpertusa Brodo

Pertusaria tetrathalamia (Fée) Nyl.

Pertusaria texana Müll. Arg.

+Phaeocalicium polyporaeum (Nyl.) Tibell

Phaeographis brasiliensis (A. Massal.) Kalb & Matthes-Leicht

Phaeographis erumpens (Nyl.) Müll. Arg.

Phaeographis inusta (Ach.) Müll. Arg.

Phaeographis lobata (Eschw.) Müll. Arg.

Phaeographis oricola Lendemer & R.C. Harris

Phaeophyscia pusilloides (Zahlbr.) Essl.

Phaeophyscia rubropulchra (Degel.) Essl.

Phaeophyscia squarrosa Kashiw.

*Phaeosporobolus alpinus R. Sant., Alstrup, & D. Hawksw.

Phlyctis boliviensis Nyl.

Phyllopsora confusa Swinsc. & Krog

Phyllopsora parvifolia (Pers.) Müll. Arg.

Physcia americana G. Merr.

Physcia atrostriata Moberg

Physcia millegrana Degel.

Physcia pumilior R.C. Harris

Physcia sorediosa (Vain.) Lynge

Piccolia nannaria (Tuck.) Lendemer & Beeching

Placynthiella dasaea (Stirt.) Tønsberg

Placynthiella icmalea (Ach.) Coppins & P. James

Polymeridium proponens (Nyl.) R.C. Harris

Polymeridium quinqueseptatum (Nyl.) R.C. Harris

Polymeridium subcinereum (Nyl.) R.C. Harris

Porina heterospora (Fink) R.C. Harris

Porina scabrida R.C. Harris

*Pronectria subimperspicua (Speg.) Lowen

Protoparmelia isidiata Diederich, Aptroot & Sérus.

Pseudosagedia cestrensis (Tuck.) R.C. Harris

Pseudosagedia isidiata (R.C. Harris) R.C. Harris

Pseudosagedia rhaphidosperma (Müll. Arg.) R.C. Harris

Psoroglaena dictyospora (Orange) H. Harada

Punctelia missouriensis G. Wilh. & Ladd

Punctelia rudecta (Ach.) Krog

Pyrenula anomala (Ach.) R.C. Harris

Pyrenula aspistea Ach.

Pyrenula citriformis R.C. Harris

Pyrenula cruenta (Mont.) Vain.

Pyrenula leucostoma Ach.

Pyrenula mamillana (Ach.) Trevis.

Pyrenula microcarpa Müll. Arg.

Pyrenula microtheca R.C. Harris

Pyrenula pseudobufonia (Rehm) R.C. Harris

Pyrenula punctella (Nyl.) Trevis.

Pyrenula ravenelii (Tuck.) R.C. Harris

Pyrenula santensis (Nyl.) Müll. Arg.

Pyrgillus javanicus Nyl.

Pyrrhospora sp. This species, widespread in the southeastern Coastal Plain, has been confused with Pyrrhospora quernea (Dicks.) Körb. when collected. It differs from that species in several respects and will be described in a future publication.

Pyrrhospora varians (Ach.) R.C. Harris

Pyxine albovirens (G. Mey.) Aptroot

Pyxine caesiopruinosa (Nyl.) Imshaug

Pyxine sorediata (Ach.) Mont.

Pyxine subcinerea Stirt.

Ramalina complanata (Sw.) Ach.

Ramalina culbersoniorum LaGreca

Ramalina stenospora Müll. Arg.

Ramalina willeyi R. Howe

Ramboldia russula (Ach.) Kalb, Lumbsch & Elix

Ramonia microspora Vězda

Rinodina dolichospora Malme

Rinodina maculans Müll. Arg.

Rinodina papillata H. Magn.

Ropalospora viridis (Tønsberg) Tønsberg

*Roselliniopsis tropica Matzer & R. Sant.

Sarcographa tricosa (Ach.) Müll. Arg.

Schismatomma rappii (Zahlbr.) R.C. Harris

Schrakia sp.? This is a very unusual species that does not appear to be lichenized, but nonetheless would easily be confused with members of the genus Melaspilea on account of its brown, two-celled ascospores. It typically occurs on the bark of trees in swamp forests and can easily be recognized in the field by the presence of small, brownish-black apothecia with distinctly red pruinose margins. We include the species here because it is not uncommon and we have been unable to locate a name for it.

Segestria leptalea (Durieu & Mont.) R.C. Harris

*Skyttea lecanorae Diederich & Etayo

*Sphinctrina tubiformis A. Massal.

Sticta carolinensis T. McDonald

Sticta deyana Lendemer & Goffinet

Strigula americana R.C. Harris

Strigula viridiseda (Nyl.) R.C. Harris

*Taeniolella delicata M.S. Christ. & D. Hawksw.

Teloschistes chrysophthalmus (L.) Tuck.

Tephromela atra (Huds.) Hafellner

Thalloloma cf. cinnabarinum (Fée) Staiger

Thalloloma hypoleptum (Nyl.) Staiger

Thelopsis rubella Nyl.

Thelotrema adjectum Nyl.

Thelotrema defectum R.C. Harris

Thelotrema dilatatum (Müll. Arg.) Hale

Thelotrema lathraeum Tuck.

Thelotrema monospermum R.C. Harris

Thelotrema subtile Tuck.

Topelia aperiens P. M. Jørg. & Vězda

Trapeliopsis flexuosa (Fr.) Coppins & P. James

*Tremella parmeliarum Diederich

*Tremella harrisii Diederich

*Tremella pertusariae Diederich

*Tremella phaeographidis Diederich, Coppins & Bandoni

*Tremella sp. This taxon was found growing on a thallus of Lecanora louisianae. Because only one small specimen was available, we have refrained from studying it further until additional material is located.

*Trichosphaerella buckii R.C. Harris & Lendemer

Trichothelium americanum Lendemer

Trypethelium tropicum (Ach.) Müll. Arg.

Trypethelium virens Tuck.

Tuckermanella fendleri (Nyl.) Essl.

Usnea baileyi (Stirt.) Zahlbr.

Usnea endochrysea Stirt.

Usnea evansii Motyka

Usnea mutabilis Stirt.

Usnea pensylvanica Motyka

Usnea strigosa (Ach.) Eaton

Usnea subscabrosa Motyka

Usnea trichodea Ach.

Variolaria amara Ach.

Variolaria commutata (Mull. Arg.) ined. (≡Pertusria commutata Mull. Arg.)

Variolaria hypothamnolica (Dibben) ined. (≡Pertusria hypothamnolica Dibben)

Variolaria multipunctoides (Dibben) Lendemer, B.P. Hodk., & R.C. Harris

Variolaria ophthalmiza (Nyl.) Darb.

Variolaria pustulata (Brodo & W.L. Culb.) Lendemer, B.P. Hodk., & R.C. Harris

Variolaria trachythallina (Erichsen) Lendemer, B.P. Hodk., & R.C. Harris

Vezdaea leprosa (P. James) Vězda

*Vouauxiella lichenicola (Linds.) Petr. & Syd.

Xyleborus nigricans R.C. Harris & Lendemer


Below we present keys to the lichens, lichenicolous fungi and allied fungi that occur in the DRBH. The keys are arranged with a main “Key to Keys” followed by subsequent smaller keys. Note that taxa included in brackets have not yet been found in the study area, but are included either to broaden the use of the keys or because there is a high chance that they occur in the DRBH. Terminology has been simplified as much as possible. We suggest that the reader refer to the introductory material and glossary of Brodo et al. (2001) for questions relating to the meaning of a given technical term.


1. Fungi occurring on the thalli of other lichensKey 1. Lichenicolous Fungi

1. Fungi not on the thalli of other lichens2

2. Fruiting body resembling white, fleshy clubs; fungus a basidiomycete Multiclavula mucida (Fr.) R. H. Petersen

2. Fruiting body not resembling white, fleshy clubs; fungus not a basidiomycete3

3. Ascomata resembling tiny black or brown pinsKey 2. Calicioid Fungi

3. Ascomata not as above4

4. Thallus foliose, fruticose or squamulose (macrolichens)5

5. Thallus foliose6

6. Photobiont a cyanobacterium, sometimes restricted to cephalodia with the primary photobiont being a green alga (all species rare or extirpated) Key 3. Foliose Cyanolichens

6. Photobiont a green alga, no cyanobacteria presentKey 4. Foliose Chlorolichens

5. Thallus fruticose or large squamulose7

7. Thallus large squamulose or dimorphic with primary thallus of squamules and secondary thallus of hollow podetia; Cladonia/CladinaKey 5. Cladoniaceae

7. Thallus not large squamulose or dimorphic, always fruticose; not Cladonia/CladinaKey 6. Fruticose Macrolichens

4. Thallus crustose8

8. Thallus on leavesKey 7. Foliicolous Lichens

8. Thallus on other substrates9

9. Thallus with lichenized diaspores (e.g., isidia, soredia) or specialized conidium bearing structures (i.e., hyphophores or stalked pycnidia), apothecia/perithecia not typically presentKey 8. Typically Asexually Reproducing Crustose Lichens

9. Thallus without lichenized diaspores or specialized conidium bearing structures10

10. Fruiting body a peritheciumKey 9. Crustose Pyrenolichens

10. Fruiting body an apothecium11

11. Ascospores colorless Key 10. Crustose Apotheciate Lichens with Hyaline Spores

11. Ascospores brown Key 11. Crustose Apotheciate Lichens with Brown Spores


1. Spores produced in asci in disciform, lirelliform or flask shaped structures2

2. Ascomata disk-like (apothecioid) or lirelliform (if hemispherical apothecioid but without asci, see Tremella)3

3. Spores soon brown, simple or 1-septate4

4. Ascomata with short stalk; spores simple, citriform (lemon-shaped), coarsely ridged Sphinctrina tubiformis A. Massal

4. Ascomata not stalked; spores 1-septate5

5. Ascomata lirelliform; on Pyrenula cruenta Melanographa tribulodes (Tuck.) Müll. Arg.

5. Ascomata apothecioid; on other hosts6

6. On thallus of Parmotrema subrigidum; epihymenium K+ green Abrothallus parmotrematis Diederich

6. On warts and thallus of Pertusaria or Bathelium carolinianum7

7. On Bathelium carolinianum; spores 16–19 × 8–11.5 μm Buelliella trypethelii (Tuck.) Fink

7. On Pertusaria8

8. Asci with I+ blue cap; spores 12–15 × 6–8.5 μm; on Pertusaria paratuberculiferaDactylospora inquilina (Tuck.) Hafellner

8. Asci without I+ cap; spores 15–18 × 6–8 μm on Pertusaria tetrathalamia Buelliella minimula (Tuck.) Fink

3. Spores colorless (may become brown with age), simple to 3-septate9

9. Ascomata dark, K−10

10. Spores simple11

11. On thallus of Lecanora floridula and L. louisianae; ascomata with marginal hairs; asci 8-spored, ellipsoid, 7–9 × 3–3.5 μmSkyttea lecanorae Diederich & Etayo

11. On thallus of Parmotrema submarginale & P. subrigidum; ascomata without marginal hairs; asci multisporous; spores ± globose to broadly ellipsoid Gyalideopsis floridae Etayo & Diederich

10. Spores 1-3-septate12

12. Spores 1-septate13

13. Ascomata not stalked, not mazaedial; on Haematomma accolens or Pyrenula cruenta14

14. Growing in hymenium of Haematomma accolens; ascomata blotchy, irregularly sharpedArthonia stevensoniana R.C. Harris & Lendemer

14. Growing on thallus surface of Pyrenula cruenta; ascomata lirelliform Melanographa tribulodes (Tuck.) Müll. Arg.

13. Ascomata with short stalk, mazaedial; on apothecia and thallus of Lecanora caesiorubella ssp. glaucomodes, L. louisianae and L. subpallens [Chaenothecopsis kalbii Tibell & K. Ryman]

12. Spores 2-3-septate15

15. On thallus of Lecanora louisianae; spores 2(−3)-septate, 13–16 × 5–7.5 μm Arthonia agelastica R.C. Harris & Lendemer

15. On Pertusaria; spores 3-septate, 17–26 × 6.5–9 μm Opegrapha anomaea Nyl.

9. Ascomata orange, KOH+ purple or whitish to pinkish16

16. Apothecia orange, K+ red; spores 3-septate, macrocephalic, 12–15 × 4–7 μm; on Graphis lineolaArthonia hodgesii Lendemer & R.C. Harris

16. Apothecia whitish to pinkish; margin concolorous or paler than disk; epihymenium and exciple filled with small POL+ crystals; spores needle shaped; over thallus of Trypethelium tropicumBacidia crystallifera Ekman

2. Ascomata flask shaped (perithecioid); interascal hyphae absent or indistinct17

17. Spores brown18

18. Asci multisporous; spores 1-septate; on various crustose lichens (incl. Buellia curtisii, Lecanora strobilina, Pertusaria epixantha and P. obruta) Muellerella lichenicola (Sommerf.) D. Hawksw.

18. Asci 8-spored19

19. Ascospores simple; on apothecia and thallus of Ochrolechia africana Roselliniopsis tropica Matzer & R. Sant.

19. Ascospores submuriform; on Rinodina maculansMerismatium sp.

17. Spores colorless or pale brownish (may turn dark brown in age), 1-septate20

20. Perithecia pink, orangish or reddish21

21. Perithecia immersed in thallus of Punctelia rudecta; wall orangish, K+ purplish spores uniseriate, 8–10 × 5–6 μm, wartyPronectria subimperspicua (Speg.) Lowen

21. Perithecia superficial; pink or reddish; wall K−22

22. Perithecia pink, fuzzy; spores of two sizes [Ovicuculospora parmeliae (Berk. & M. A. Curtis) Etayo]

22. Perithecia reddish with short colorless setae; spores all one size; on various crustose lichens (incl. Micarea prasina) Nectriopsis rubefaciens (Ellis & Everh.) M.S. Cole & D. Hawksw.

20. Perithecia black23

23. Asci 8-spored; on Haematomma persoonii Lichenochora haematommatum R.C. Harris & Lendemer

23. Asci 16-spored (part spores); part spores tetrahedral; perithecia with setae around ostiole; in thallus of Punctelia rudecta Trichosphaerella buckii R.C. Harris & Lendemer

1. Spores not produced in asci, but rather in flask-shaped or disciform structures (pycnidia, sporodochia), tiny stromata (Phaeosporobolus) or basidiomata (Tremella) or from hyphae embedded in thallus or apothecia (“hyphomycetes”)24

24. Spores produced in pycnidia, sporodochia, or tiny stromata25

25. Spores brown or greenish26

26. Conidioma a pycnidium27

27. Conidia 1-septate28

28. Conidia formed in short chains, greenish; on apothecia and thallus of Lecanora louisianae, L. subpallens and Pyrrospora varians Vouauxiella lichenicola Petr. & Syd.

28. Conidia not formed in chains, brown; on Pertusaria. Lichenodiplis lecanorae (Vouaux) Dyko & D. Hawksw.

27. Conidia simple29

29. Conidia irregularly shaped, 8–13 × 5–6 μm; on Leiorreuma and Phaeographis Coniambigua phaeographidis Etayo & Diederich

29. Conidia ± globose to pyriform30

30. Conidia globose, 5–6 × 4.5–5 μm; on Lecanora hybocarpa Lichenoconium lecanorae (Jaap) D. Hawksw.

30. Conidia pyriform, 8–10 × 3–3.5 μm; on Parmotrema subrigidum Lichenoconium cargillianm (Linds.) D. Hawksw.

26. Conidioma a sporodochium or tiny superficial stroma31

31. Conidia produced in sporodochia32

32. Exciple orange red; conidia simple, 4.7–5.8 × 3–4 μm; on Pertusaria epixanthaMinutoexcipula miniatoexcipula R.C. Harris & Lendemer

32. Exciple brown; conidia 1-septate33

33. Conidia 6.5–8 × 3–4 μm; on Pertusaria texana Minutoexcipula tuckerae V. Atienza & D. Hawksw.

33. Conidia 6–7.5 × 2.5–3.5 μm; on Pertusaria pustulata and P. subpertusa Minutoexcipula mariana V. Atienza

31. Conidia produced in tiny orbicular or slightly irregular stromata; conidia muriform, ± globose34

34. On various crustose lichens, especially Phaeographis and Leiorreuma Etayoa trypethelii (Flakus & Kukwa) Diederich & Ertz

34. On Ochrolecia and Pertusaria.Phaeosporobolus alpinus R. Sant. et al.

25. Spores colorless35

35. Conidia broadly ellipsoid, 3.5–6 × 3–4 μm, with a single guttule; on Parmeliaceae, esp. Punctelia rudectaDidymocyrtis melanelixiae (Brackel) Diederich, R.C. Harris & Etayo (Phoma asexual stage)

35. Conidia tetrahedral (triangular in surface view); apices papillate on Pertusaria subpertusaCornutispora triangularis Diederich & Etayo

24. Spores produced by hyphae embedded in apothecia or thallus (“hyphomycetes”) or basidiomata (not in pycnidia or sporodochia)36

36. Hyphomycetes37

37. Conidia 1(−2)-septate, 7–11 × 3.5–6 μm; on Lecanora louisianae and L. subpallens Taeniolella delicata M.S. Christ. & D. Hawksw.

37. Conidia 0-septate, 3–4.5 × 2.5–4 μm, forming chains; in hymenium of Byssoloma meadii Intralichen lichenum (Diederich) D. Hawksw. & M.S. Cole

36. Basidiomycetes38

38. On Parmotrema perforatumTremella parmeliarum Diederich

38. On PertusariaTremella pertusariae Diederich

38. On LeiorreumaTremella phaeographidis Diederich, Coppins & Bandoni

38. On Bathelium carolinianumTremella harrisii Diederich

38. On Lecanora louisianaeTremella sp.


1. Ascospores simple2

2. Ascospores ellipsoid; thallus not evident; on bark and woodMycocalicium subtile (Pers.) Szatala

2. Ascospores globose, simple; thallus granulose, green; on Chamaecyparis and Taxodium Chaenotheca hygrophila Tibell

1. Ascospores 2-celled3

3. On the polypore Trichaptum biformePhaeocalicium polyporaeum (Nyl.) Tibell

3. On wood4

4. Capitulum K+ redChaenothecopsis pusiola (Ach.) Vain.

4. Capitulum K- or K+ intensifying, but not K+ red5

5. Ascospores with septum distinctly more lightly pigmented than the walls Chaenothecopsis pusilla (Ach.) A.F.W. Schmidt

5. Ascospores with normally pigmented septum just as dark as the walls6

6. Stalks tall, reddish pigmented, C+ fleeting greenish in section Chaenothecopsis debilis (Sm.) Tibell

6. Stalks short, not reddish pigmented, C-Chaenothecopsis nana Tibell


1. Thallus isidiate or lobulate, with lichenized diaspores; apothecia rarely produced2

2. Medulla P+ orange-red, isidiatePannaria tavaresii P.M. Jørg.

2. Medulla P-, isidiate or lobulate3

3. Thallus slate or lead gray4

4. Thallus thick, with a conspicuous weft of rhizines not gelatinous when wet5

5. Thallus isidiate; commonCoccocarpia palmicola (Spreng.) Arv. & D.J. Galloway

5. Thallus lobulate; rareParmeliella pannosa (Sw.) Müll. Arg.

4. Thallus thin, without rhizines, gelatinous when wet6

6. Thallus smooth, isidia or lobules distributed more-or-less evenly across the surface Leptogium cyanescens (Rabenh.) Körb.

6. Thallus wrinkles, isidia or lobules often somewhat concentrated on the wrinkled Leptogium austroamericanum (Malme) C.W. Dodge

3. Thallus black or brown7

7. Thallus black; isidia globoseCollema subflaccidum Degel.

7. Thallus brown; isidiate or lobulate8

8. Thallus fruticose, a complex tangle of intensely divided minute lobes, resembling Leptogium lichenoidesDendriscocaulon intricatulum (Nyl.) Henssen

8. Thallus foliose, with distinct plane lobes bearing marginal phyllidia9

9. Medulla white, K-Sticta carolinensis T. McDonald

9. Medulla orange, K+ purpleSticta deyana Lendemer & Goffinet

1. Thallus without lichenized diaspores; apothecia frequently produced10

10. Thallus growing on the ground or over organic matter at the bases of trees; lower surface with a distinct network of veins and long rhizinesPeltigera neopolydactyla (Gyeln.) Gyeln.

10. Thallus corticolous; lower surface not as above11

11. Thallus black, ridged and pustulose; apothecia densely white pruinose Collema pulcellum Ach. var. leucopeplum (Tuck.) Degel.

11. Thallus varying shades of gray, not ridged and pustulose; apothecia not densely white pruinose12

12. Thallus composed of overlapping squamules, without distinct marginal radiating lobes Fuscopannaria leucosticta (Tuck.) P. M. Jørg.

12. Thallus truly foliose, with distinct marginal lobes13

13. Thallus gelatinous (jelly-like) when wet14

14. Lower surface with abundant rhizines Pannaria lurida (Mont.) Nyl. subsp. quercicola P. M. Jørg.

14. Lower surface without rhizines15

15. Thallus smoothLeptogium azureum (Sw.) Mont.

15. Thallus wrinkledLeptogium corticola (Taylor) Tuck.

13. Thallus not gelatinous (not jelly-like) when wet16

16. Thallus lead gray; with a distinct weft of dark rhizines; main photobiont a cyanobacteriumCoccocarpia erythroxyli (Spreng.) Swinsc. & Krog

16. Thallus gray to greenish; with pale rhizines if rhizines are present; main photobiont a green algaLobaria ravenelii (Tuck.) Yoshim.


1. Thallus not pustulose, sorediate, isidiate, or phyllidiate; apothecia often present2

2. Thallus with a distinct black, felt-like hypothallus on the lower surface; very rare Anzia colpodes (Ach.) Stizenb.

2. Thallus with a naked lower surface or with rhizines, but not with a distinct black, felt-like hypothallus; common3

3. Lobes broad, >3 mm wide; ascospores hyaline4

4. Lobes adnate; thallus wrinkled, scrobiculate upper surface; medulla C+ pink (gyrophoric acid present); rareLobaria ravenelii (Tuck.) Yoshim.

4. Lobes adnate or ascending; thallus with a smooth upper surface; medulla C- other substances present)5

5. Apothecia entire, not perforated with a hole in the center; medulla P+ red (protocetraric acid present)Parmotrema submarginale (Michx.) DePriest & B. Hale

5. Apothecia perforated with a hole in the center; medulla P- or P+ yellow (alectoronic and/or norstictic acid present)6

6. Medulla UV- (alectoronic acid absent); typically inland Parmotrema perforatum (Jacq.) A. Massal.

6. Medulla UV+ bright blue-white (alectoronic acid present); typically coastal, rarer inland7

7. Medulla, especially near the apothecia, K- (norstictic acid absent) Parmotrema subrigidum Egan s. str.

7. Medulla, especially near the apothecia, K+ yellow turning red (norstictic acid present)Parmotrema subrigidum Egan s. lat.

3. Lobes narrow, <3 mm wide; ascospores brown or hyaline8

8. Thallus dark greenish-brown to brown, forming minute rosettes on conifer branches Tuckermanella fendleri (Nyl.) Essl.

8. Thallus gray to blue-gray or light brownish, not forming minute rosettes on conifer branches9

9. Medulla K+ pink turning dark brown (lividic acid present); ascospores hyaline Hypotrachyna livida (Taylor) Hale

9. Medulla K- or K+ yellow (atranorin present or absent); ascospores brown10

10. Lower surface black; medulla UV+ blue-white (sekikaic acid present); primarily coastalDirinaria confusa D.D. Awasthi

10. Lower surface white or pale brownish; medulla UV- (sekikaic acid absent); inland and coastal11

11. Upper cortex and medulla K+ yellow (atranorin present); lobes not abundantly lobulatePhyscia pumilior R.C. Harris

11. Upper cortex and medulla K- (atranorin absent); lobes abundantly lobulateAnaptychia palmulata (Michx.) Vain.

1. Thallus pustulose, sorediate, isidiate, or phyllidiate; apothecia uncommon12

12. Thallus with a distinct black, felt-like hypothallus on the lower surface; lobe margins abundantly isidiate/phyllidiateAnzia ornata (Zahlbr.) Asahina

12. Thallus with a naked lower surface or with rhizines, but not with a distinct black, felt-like hypothallus13

13. Thallus isidiate or lobulate14

14. Thallus lobulate

15. Lower surface black, at least centrally; lobules fine, erect, fragile Phaeophyscia squarrosa Kashiw.

15. Lower surface plae to tan throughout; lobules coarse and robust, not erect, not fragile Anaptychia palmulata (Michx.) Vain.

14. Thallus isidiate16

16. Lower surface ecorticate, orange pigmented; orange pigment K+ purple Heterodermia crocea R.C. Harris

16. Lower surface corticate, not orange pigmented17

17. Medulla C+ pink or red (gyrophoric or lecanoric acid present)18

18. Lobes broad, >3 mm wide19

19. Lower surface pale; upper surface with conspicuous white pseudocyphellae; conspicuous marginal cilia absent20

20. Thallus isidiate; isidia short to tall, cylindrical, brown tipped, not clustered only in the pseudocyphellae Punctelia rudecta (Ach.) Krog

20. Thallus with squamiform soredia that resemble isidia; “isidia” short, squamiform, poorly corticate, not brown tipped, always clustered in the pseudocyphellae Punctelia missouriensis G. Wilh. & Ladd

19. Lower surface black towards the center and brown towards the margin; upper surface without conspicuous white pseudocyphellae; conspicuous marginal cilia present21

21. Upper surface yellow-green, K-, KC+ strong yellow (usnic acid present)Parmotrema madagascariaceum (Hue) Hale

21. Upper surface blue-gray, K+ yellow, KC- (atranorin present) Parmotrema tinctorum (Nyl.) Hale

18. Lobes narrow, <3 mm wide22

22. Marginal cilia with bulbate bases; lower surface brown Bulbothrix scortella (Nyl.) Hale

22. Marginal cilia without bulbate bases; lower surface black Hypotrachyna minarum (Vain.) Krog & Swinsc.

17. Medulla C- (gyrophoric or lecanoric acid absent)23

23. Medulla P+ orange or red (protocetraric acid, salazinic acid or stictic acid present)24

24. Marginal cilia present, with bulbate bases; lobes narrow, <3 mm wide Bulbothrix isidiza (Nyl.) Hale

24. Marginal cilia present or absent, but always lacking bulbate bases; lobes broad, >3 mm wide25

25. Medulla K+ yellow turning dirty brown (protocetraric or stictic acid present); marginal cilia absent to sparse26

26. Medulla P+ orange (stictic acid present together with norlobaridone); upper surface without pseudocyphellae Parmotrema internexum (Nyl.) Hale

26. Medulla P+ red (protocetraric acid present); upper surface with white pseudocyphellae Canoparmelia amazonica (Nyl.) Elix & Hale

25. Medulla K+ yellow turning red (salazinic acid present); marginal cilia usually conspicuously present and abundant27

27. Lower portions of medulla UV+ bright yellow (lichexanthone present)Parmotrema ultralucens (Krog) Hale

27. Entire medulla UV-28

28. Lower surface black Parmotrema subisidiosum (Müll. Arg.) Hale

28. Lower surface brown Parmotrema neotropicum Kurok.

23. Medulla P- (salazinic acid and stictic acid absent)29

29. Lobes broad, >3 mm wide30

30. Lobes adnate; margins without long conspicuous cilia Canoparmelia caroliniana (Nyl.) Elix & Hale

30. Lobes ascending; marginal with long conspicuous cilia31

31. Medulla UV+ blue-white (alectoronic acid present); upper cortex blue-gray, K+ yellow, KC- (atranorin present) Parmotrema mellissii (C.W. Dodge) Hale

31. Medulla UV- (fatty acids present); upper cortex yellow-green, K-, KC+ yellow gold (usnic acid present) Parmotrema xanthinum (Müll. Arg.) Hale

29. Lobes narrower, <3 mm wide32

32. Isidia never ciliate, often breaking down into piles that resemble soralia with large coarse soredia; lobes tightly adnate; medulla UV+ blue-white (divaricatic acid present); primarily coastal in the DRBHDirinaria aegialita (Ach.) B.J. Moore

32. Isidia ciliate, never breaking down into soralia; lobes adnate, but not tightly so; medulla UV- (divaricatic acid absent); throughout the DRBHHypotrachyna horrescens (Taylor) Swinsc. & Krog

13. Thallus pustulose or sorediate33

33. Upper cortex UV+ bright yellow (lichexanthone present)34

34. Medulla white; lower surface with dichotomously branched and forking rhizines Hypotrachyna osseoalba (Vain.) Y.S. Park & Hale

34. Medulla yellow or orange pigmented; lower surface with simple or forking rhizines35

35. Lobes with dactyls along the marginsPyxine caesiopruinosa (Nyl.) Imshaug

35. Lobes with discrete, laminal and marginal soralia36

36. Lobe tips typically with discrete white “pads” of pruina; medulla K- Pyxine subcinerea Stirt.

36. Lobe tips without discrete white “pads” of pruina; medulla K+ purple (but often difficult to detect)Pyxine albovirens (G. Mey.) Aptroot

33. Upper cortex UV- (lichexanthone absent)37

37. Lower surface ecorticate38

38. Lower surface yellow or orange pigmented, at least in spots near the lobe tips39

39. Lower surface orange pigmented; pigment K+ purple Heterodermia obscurata (Nyl.) Trevis.

39. Lower surface with spots of yellow pigment near the lobe tips; pigment K-Heterodermia casarettiana (A. Massal.) Trevis.

38. Lower surface not yellow or orange pigmented40

40. Lobes elongate, linear, strap-shaped, +/- ascending Heterodermia leucomelos (L.) Poelt

40. Lobes not linear and strap-shaped, always adnate41

41. Lower surface weakly corticate; upper surface shiny, not appearing frosted; medulla K+ yellow turning red but without norstictic acid crystals (salazinic acid present) Heterodermia albicans (Pers.) Swinsc. & Krog

41. Lower surface entirely ecorticate; upper surface appearing frosted with a white pruina; medulla K+ yellow (salazinic acid absent) Physcia atrostriata Moberg

37. Lower surface corticate42

42. Medulla orange-red or yellow pigmented43

43. Medulla strongly orange-red pigmented; pigment K+ purple Phaeophyscia rubropulchra (Degel.) Essl.

43. Medulla yellow pigmented; pigment K-44

44. Thallus with discrete soralia; medulla strongly yellow pigmented Pyxine sorediata (Ach.) Mont.

44. Thallus with laminal or marginal pustules; medulla weakly yellow pigmented45

45. Medulla C+ pink (gyrophoric acid present); thallus with coarse pustulesHypotrachyna spumosa (Asahina) Krog & Swinsc.

45. Medulla C- (gyrophoric acid absent); thallus with diffuse pustules Myelochroa aurulenta (Tuck.) Elix & Hale

42. Medulla white46

46. Upper surface yellow or yellow green (calycin or usnic acid present)47

47. Soralia marginal/terminal, not laminal; upper surface yellow, KC- (calcyin present)Candelaria concolor (Dicks.) Stein

47. Soralia laminal; upper surface yellow-green, KC+ yellow (usnic acid present)48

48. Lobes broad, >3 mm wide; medulla P+ orange-red, UV- (protocetraric acid present)Flavoparmelia caperata (L.) Hale

48. Lobes narrow, <1 mm wide; medulla P-, UV+ blue-white (divaricatic acid present)Parmeliopsis subambigua Gyeln.

46. Upper surface blue-gray or gray-brown (atranorin present or absent)49

49. Upper cortex gray-brown, K- (atranorin absent); capitate soralia terminal on the tips of secondary lobes Phaeophyscia pusilloides (Zahlbr.) Essl.

49. Upper cortex blue-gray, K+ yellow (atranorin present); soralia not as above50

50. Medulla C+ pink-red (gyrophoric or lecanoric acid present)51

51. Lower surface pale; upper surface with conspicuous white pseudocyphellae; squamiform soredioid-isidia present and clustered in the pseudocyphellae Punctelia missouriensis G. Wilh. & Ladd

51. Lower surface black; upper surface without conspicuous white pseudocyphellae; soralia or pustules present52

52. Thallus with discrete soralia; rare Hypotrachyna cryptochlora (Vain.) D. Hawksw. & A. Crespo

52. Thallus with coarse pustules; common Hypotrachyna spumosa (Asahina) Krog & Swinsc.

50. Medulla C- (gyrophoric or lecanoric acid absent)53

53. Medulla K+ yellow turning red (salazinic acid or norstictic acid present)54

54. Lobes narrow, <3 mm wide; black marginal cilia absent Heterodermia albicans (Pers.) Swinsc. & Krog

54. Lobes broad, >3 mm wide; black marginal cilia present55

55. Lobes strongly ascending; lower surface with broad white blotches especially near the margins56

56. Medulla K+ yellow turning red, producing norstictic acid crystals, P+ yellow (norstictic acid present) Parmotrema hypotropum (Nyl.) Hale

56. Medulla K+ yellow turning red, not producing norstictic acid crystal, P+ orange (stictic acid present)Parmotrema hypoleucinum (J. Steiner) Hale

55. Lobes not strongly ascending; lower surface entirely black in the center and brown towards the margins57

57. Capitate soralia with fine soredia present on the tips of the sublobes Parmotrema reticulatum (Taylor) M. Choisy

57. Diffuse soralia with coarse soredia present on the tips of the sublobes and lobes as well as on the thallus surface Parmotrema subisidiosum (Müll. Arg.) Hale

53. Medulla K- or K+ yellow-brown, but not K+ yellow turning red58

58. Medulla P+ orange or red (protocetraric acid or stictic acid present)59

59. Medulla P+ orange (stictic acid present); thallus surface scrobiculateCrespoa crozalsiana (Harm.) Lendemer & B.P. Hodk.

59. Medulla P+ red (protocetraric acid present); thallus surface smooth, not scrobiculate60

60. Echinocarpic acid absent; common [TLC required] Parmotrema gardneri (C.W. Dodge) Sérus.

60. Echinocarpic acid present; rare [TLC required] Parmotrema dilatatum (Vain.) Hale

58. Medulla P- (protocetraric acid or stictic acid absent)61

62. Lobes broad, >3 mm wide63

63. Medulla UV- (fatty acids present) Parmotrema praesorediosum (Nyl.) Hale

63. Medulla UV+ blue-white (alectoronic acid present)64

64. Thallus with marginal soralia Parmotrema rampoddense (Nyl.) Hale

64. Thallus with coarse, laminal soredia that arise from the breakdown of isidia Parmotrema mellissii (C.W. Dodge) Hale

62. Lobes narrow, <3 mm wide65

65. Medulla UV+ blue-white (divaricatic acid present) Dirinaria picta (Sw.) Schaer. ex Clem.

65. Medulla UV- (divaricatic acid present)66

66. Lower surface black Physcia sorediosa (Vain.) Lynge

66. Lower surface pale67

67. Thallus with discrete, laminal soralia; upper surface often pruinose; common Physcia americana G. Merr.

67. Thallus with continuous, marginal soralia; upper surface epruinose; rare Physcia millegrana Degel.


1. Podetia present; thallus with or without primary squamules2

2. Podetia abundantly branching and forming cushions on the ground3

3. Podetia corticate; apothecia/pycnidia redCladonia leporina Fr.

3. Podetia ecorticate; apothecia/pycnidia pallid to brown or unknown4

4. Podetia (especially near the tips) P-, UV+ blue-white (perlatolic acid present); restricted to barrier islands in the DRBHCladonia evansii Abbayes

4. Podetia (especially near the tips) P+ red, UV- (fumarprotocetraric acid present); common on roadsides and sandy soils throughout the DRBHCladonia subtenuis (Abbayes) Mattick

2. Podetia simple or little branching, but not forming cushions on the ground5

5. Podetia with a continuous cortex, not sorediate or microsquamulose6

6. Podetia forming multi-tiered cups that proliferate from the centers, resembling a wedding cakeCladonia rappii A. Evans

6. Podetia not forming cups7

7. Apothecia/pycnidia red; primary squamules conspicuously sorediate; thallus P- Cladonia incrassata Flörke

7. Apothecia/pycnidia brown; primary squamules esorediate; thallus P+ yellow, orange or red8

8. Podetia tall, little branching, forming funnels; thallus UV+ blue-white and P+ yellow (baeomycesic acid and squamatic acid present) Cladonia atlantica A. Evans

8. Podetia short, blunt, not forming funnels; thallus UV- or UV+ dull white, P+ orange or red (fumarprotocetraric acid or stictic acid present)9

9. Podetia slender, conspicuously overtopped by large brown apothecia; primary squamules small, not lobed, decumbent and overlapping; fumarprotocetraric acid present Cladonia peziziformis (With.) J.R. Laundon

9. Podetia broad, not conspicuously overtopped by the brown apothecia; primary squamules large, lobed, erect; atranorin, norstictic acid and stictic acid presentCladonia polycarpia G. Merr.

5. Podetia with a discontinuous cortex, with some parts dissolving into soredia or microsquamules10

10. Apothecia/pycnidia red11

11. Podetia sorediate, with the soredia abrading to reveal remnants of the loose medulla or the opaque white stereome; thallus P- (barbatic acid present) Cladonia macilenta var. bacillaris (Genth) Schaer.

11. Podetia microsquamulose, with the microsquamules sloughing off to reveal the naked translucent stereome; thallus P- (barbatic acid present) or P+ orange (thamnolic acid present)12

12. Thallus P+ orange (thamnolic acid present) Cladonia didyma var. vulcanica (Zoll. & Moritzi) Vain.

12. Thallus P- (barbatic acid present).Cladonia didyma var. didyma (Fée) Vain.

10. Apothecia/pycnidia brown13

13. Thallus UV+ blue-white and P+ yellow (squamatic acid and baeomycesic acid present) Cladonia beaumontii (Tuck.) Vain.

13. Thallus UV- and P+ orange or red (other substances present)14

14. Thallus K+ instantly lemon yellow, P+ orange (thamnolic acid present)15

15. Squamules small, +/- erect, dissolving into “isidioid” microsquamules; podetia slender, covered with microsquamules, the microsquamules sloughing off to reveal the naked translucent stereome Cladonia parasitica (Hoffm.) Hoffm.

15. Squamules large, decumbent, not dissolving into microsquamules; podetia short, broad, covered with coarse squamules that are not easily dislodgedCladonia santensis Tuck.

14. Thallus K- or K+ dingy yellow-brown, P+ red (fumarprotocetraric acid present)16

16. Primary squamules small, not distinctly lobed, entirely dissolving into soredia; podetia often deformed and tortuous Cladonia ramulosa (With.) J.R. Laundon

16. Primary squamules large, lobed, not entirely dissolving into soredia; podetia blunt, never deformed and tortuous17

17. Podetia with the basal portions remaining corticate, often with sorediate patches in the lower portions admixed with intact corticate areas; microsquamules absentCladonia ochrochlora Flörke

17. Podetia with the basal portions microsquamulose, often with the upper and middle portions coarsely sorediate; microsquamules present Cladonia subradiata (Vain.) Sandst.

1. Podetia absent; thallus entirely composed of primary squamules18

18. Conspicuous apothecia present, borne directly on a short stipe arising from the primary squamules Cladonia caespiticia (Pers.) Flörke

18. Apothecia absent19

19. Thallus P-; pycnidia red20

20. Primary squamules large, broad, little lobed, conspicuously sorediate; squamatic acid present Cladonia incrassata Flörke

20. Primary squamules small, lobes, not conspicuously sorediate; barbatic acid present Cladonia macilenta var. bacillaris (Genth) Schaer.

19. Thallus P+ yellow, orange or red; pycnidia brown or red21

21. Thallus P+ yellow (baeomycesic acid present)22

22. On sandy soil in disturbed areasCladonia atlantica A. Evans

22. On bark or rotting wood in swampsCladonia beaumontii (Tuck.) Vain.

21. Thallus P+ orange or red (fumarprotocetraric acid, stictic acid, or thamnolic acid present)23

23. Thallus K+ instantly lemon yellow (thamnolic acid present)24

24. Primary squamules robust, broad, little lobed, not dissolving Cladonia santensis Tuck.

24. Primary squamules fragile, narrow, lobed, often dissolving25

25. Primary squamules often entirely dissolving into “isidioid” microsquamules; pycnidia brownCladonia parasitica (Hoffm.) Hoffm.

25. Primary squamules typically remaining intact, sometimes dissolving into soredia; pycnidia red Cladonia didyma var. vulcanica (Zoll. & Moritzi) Vain.

23. Thallus K- or K+ dingy yellow-brown26

26. Thallus P+ orange (stictic acid present); squamules erect, not dissolving Cladonia polycarpia G. Merr.

26. Thallus P+ red (fumarprotocetraric acid present); squamules plane or decumbent, dissolving or not27

27. Primary squamules dissolving entirely into soredia Cladonia ramulosa (With.) J.R. Laundon

27. Primary squamules remaining intact, at least for the most part28

28. Primary squamules small, overlapping and decumbent, not lobed Cladonia peziziformis (With.) J.R. Laundon

28. Primary squamules large, not overlapping and decumbent, lobed Cladonia ochrochlora Flörke or Cladonia subradiata (Vain.) Sandst.


1. Thallus bright orange-yellow, K+ purple2

2. Thallus sorediate; apothecia rare and often absent; branches elongate, slender [Teloschistes flavicans (Sw.) Norman]

2. Thallus esorediate, apothecia often present; branches short, blade-like Teloschistes chrysophthalmus (L.) Norm.

1. Thallus gray or yellow-green, not K+ purple3

3. Branches solid without a central cord or cavity4

4. Branches smooth5

5. Branches completely flattened to the tips; perlatolic acid present Ramalina stenospora Müll. Arg.

5. Branches rounded except near the base; divaricatic or homosekikaic acid present [Ramalina montagnei De Not.]

4. Branches with abundant raised tubercles or with ridges and depressions6

6. Apothecia on the surface of the branches or along the margins Ramalina complanata (Sw.) Ach.

6. Apothecia on the tips of the branches7

7. Branches with conspicuous white tubercles; salazinic acid present or protocetraric acid presentRamalina willeyi R. Howe

7. Branched without conspicuous white tubercles; but with ridges and depressions; various substances other than the above8

8. Medulla with lichen substances present (TLC required) Ramalina culbersoniorum LaGreca

8. Medulla without lichen substances (TLC required) [Ramalina americana Hale, present just north of DRBH]

3. Branches with a central cord or cavity9

9. Thallus without soredia or isidia; apothecia typically present10

10. Restricted to coastal maritime forests; medulla white; branches often weakly foveolate or ridged; fibrils regularly arranged; galbinic acid presentUsnea evansii Motyka

10. Restricted to inland habitats; medulla white or red; branches never foveolate or ridged; fibrils haphazardly arranged; galbinic acid absent11

11. Ascospores averaging 9 μm long; medulla pink-red pigmented; norstictic acid present or absent12

12. Norstictic acid presentUsnea endochrysea Stirt. s. str.

12. Norstictic acid absentUsnea endochrysea Stirt. s. lat.

11. Ascospores averaging < 9 μm long; medulla often white, sometimes pink-red pigmented; norstictic present or absent13

13. Norstictic acid presentUsnea strigosa (Ach.) Eaton s. str.

13. Norstictic acid absent14

14. Psoromic acid presentUsnea strigosa (Ach.) Eaton s.l.

14. Psoromic acid absent15

15. Thamnolic acid presentUsnea strigosa (Ach.) Eaton s.l.

15. Thamnolic acid absentUsnea strigosa (Ach.) Eaton s.l.

9. Thallus sorediate or isidiate; apothecia rarely present16

16. Medulla pink-red pigmented17

17. Branches with a central cord; medulla P- (fatty acids present); common Usnea mutabilis Stirt.

17. Branches with a central cavity; medulla P+ yellow (norstictic acid present); rare Usnea baileyi (Stirt.) Zahlbr.

16. Medulla white18

18. Thallus long, pendant; branches with white rings (annular pseudocyphellae) Usnea trichodea Ach.

18. Thallus short, shrubby; branches without white rings19

19. Cortex red pigmented, pigment often mottledUsnea pensylvanica Motyka

19. Cortex not red pigmented20

20. Medulla P+ orange-red (protocetraric acid present) Usnea subscabrosa Motyka s. str.

20. Medulla P- (protocetraric acid absent)Usnea subscabrosa Motyka s. lat.


1. On leaves of Persea, especially in inland swamps; thallus forming tiny shiny greenish areoles, each with a central black pycnidium; only pycnidia observed; apothecia and ascospores unknown in MACP Asterothyrium decipiens (Rehm) R. Sant. ?

1. On leaves of Sabal minor or Ilex; thallus areolate or not, with either pale tan pycnidia or tall black sterile setae2

2. On leaves of Sabal minor; thallus areolate, with pale tan pycnidia; conidia bacilliform; apothecia and ascospores unknown in MACP materialFellhanera bouteillei (Desm.) Vezda

2. On leaves of Ilex; thallus continuous, with tall black sterile setae; apothecia and ascospores unknown in MACP material[Tricharia santessonii D. Hawsks.]


1. Thallus with specialized structures (e.g., hyphophores, stalked pycnidia) for the dispersal of non-lichenized diaspores (conidia); lichenized diaspores (e.g., isidia, soredia, granules) absent2

2. Thallus with hyphophores (darkened structures that resemble apically widened hairs)3

3. Hyphophores tall, erect, not apically widened, resembling an eyelash Gyalideopsis buckii Lücking et al.

3. Hyphophores short, bent down, apically widened, resembling a tiny bent shovel [Gyalideopsis ozarkensis Lücking & W. R. Buck]

2. Thallus with pale stalked pycnidia or pale synnemata (stalked sporodochia)4

4. Thallus with pale synnemata (stalked sporodochia); conidia compound aggregations of round cells [Dictyocatenulata alba Finley & E.F. Morris]

4. Thallus with pale stalked pycnidia; conidia simple, ellipsoid5

5. Thallus UV+ blue-white (lobaric acid present), often P+ orange-red (fumarprotocetraric acid present)Micarea neostipitata Coppins & P. May

5. Thallus UV- and P- (lobaric and fumarprotocetraric acid absent) [Micarea pycnidiophora Coppins & P. James]

1. Thallus without specialized structures for the dispersal of non-lichenized diaspores; lichenized diaspores present6

6. Photobiont Trentepohlia7

7. Thallus sorediate8

8. Soralia irregular in shape, diffuse; soredia light yellow-brown; thallus P-, typically indistinct and immersed in the substrateOpegrapha corticola Coppins & P. James

8. Soralia discrete, punctiform; soredia white to gray; thallus P+ orange (stictic acid present), typically distinct and white to gray in colorNadvornikia sorediata R.C. Harris

7. Thallus isidiate9

9. Thallus dark brown to grayPseudosagedia isidiata (R.C. Harris) R.C. Harris

9. Thallus yellow-brown, often bronze in color10

10. Thallus smooth and shiny; isidia typically short, +/- globose and inconspicuous; perithecia often present, dark brown-black, covered with apical setae Trichothelium americanum Lendemer

10. Thallus roughened and dull; isidia typically tall, often coralloid, conspicuous; perithecia often present, concolorous with the thallus, smooth or covered with short isidia, but never with black setaePorina scabrida R.C. Harris

6. Photobiont coccoid11

11. Thallus yellow or orange12

12. On rock (concrete); thallus orange, K+ purple[Caloplaca flavocitrina (Nyl.) H. Olivier]

12. On bark or wood; thallus yellow, K-13

13. Thallus C+ or KC+ yellow-orange (xanthones present)14

14. Thallus composed of flat green areoles with small, discrete, often marginal soralia; soredia yellow; typically on hardwoodsBuellia wheeleri R.C. Harris

14. Thallus composed of orange-yellow granules (leprose); soralia or soredia absent; typically on conifersPyrrhospora sp. nov.

13. Thallus C- and KC- (xanthones absent)15

15. Thallus areolate, with the areoles dissolving into piles of soredia; apothecia not uncommonCandelariella xanthostigmoides (Müll. Arg.) R.W. Rogers

15. Thallus leprose, composed of granules of various sizes; apothecia unknown in MACP material16

16. Granules minute (>10 μm in diameter); thallus thin, dull yellow in color (rhizocarpic acid present); on conifers in humid, swampy forests Chrysothrix chamaecyparicola Lendemer

16. Granules larger (>25–45 μm in diameter); thallus thicker, intense bright yellow in color (pinastric acid present); on conifers and hardwoods, often in upland habitatsChrysothrix xanthina (Vain.) Kalb

11. Thallus green, gray, or brown17

17. Thallus C+ pink (gyrophoric or lecanoric acid present; always confirm in squash mount if C- under the dissecting microscope)18

18. Thallus isidiatePlacynthiella icmalea (Ach.) Coppins & P. James

18. Thallus sorediate19

19. Thallus lead gray, with dark blue-gray soredia Trapeliopsis flexuosa (Fr.) Coppins & P. James

19. Thallus not lead gray, soredia various colors20

20. Thallus light gray, continuous; “soralia” regular and discoid; on hardwoods[Varicellaria velata (Turner) I. Schmitt & Lumbsch]

20. Thallus green or brown-green, areolate; soralia irregular and dissolving the areoles; on rotting wood, organic matter, or rarely the base of Pinus Placynthiella dasaea (Stirt.) Tønsberg

17. Thallus C- (gyrophoric or lecanoric acid absent)21

21. Thallus UV+ bright yellow (lichexanthone present)22

22. Thallus coarsely pustulate; medulla of pustules K- (zeorin present) Megalospora pachycheila (Tuck.) Sipman

22. Thallus with densely pruinose apothecia resembling discrete; “soralia” K+ various colors (zeorin absent)23

23. Medulla of “soralia” K+ yellow rapidly turning lavender, P- (hypothamniolic acid present)Variolaria hypothamnolica (Dibben) ined.

23. Medulla of “soralia” K+ yellow slowly turning dirty brownish or rapidly turning deep reddish-brown, P+ orange (haemathamnolic or thamnolic acid present)24

24. Medulla of “soralia” K+ yellow rapidly turning deep reddish-brown (haemathamnolic acid present); common Variolaria commutata (Mull. Arg.) ined.

24. Medulla of “soralia” K+ yellow slowly turning dirty brownish (thamnolic aicd present); rare Variolaria trachythallina (Erichsen) Lendemer et al.

21. Thallus UV- or UV+ blue-white, but not UV+ bright yellow (lichexanthone absent)25

25. Thallus K- and KC+ yellow (usnic acid present)26

26. Thallus with a well developed, fibrous, white prothallus [Lecanora thysanophora R.C. Harris]

26. Thallus without a well developed, fibrous, white prothallus27

27. Thallus areolate, with discrete circular soralia Lecanora floridula Lumbsch

27. Thallus granular to areolate, without discrete circular soralia28

28. Decarboxysquamatic acid present (TLC needed) Lecanora strobilina (Spreng.) Kieff.

28. Decarboxysquamatic acid absent (TLC needed) Lecanora cf. strobilina (Spreng.) Kieff.

25. Thallus K- or K+ yellow, always KC- (usnic acid or xanthones absent)29

29. Thallus P+ intense yellow, orange, or orange-red (fumarprotocetraric acid, pannarin, psoromic acid or thamnolic acid present)30

30. Thallus P+ intense yellow (psoromic acid present) Phlyctis boliviensis Nyl.

30. Thallus P+ orange or orange-red31

31. Thallus entirely leprose, composed of granules (Lepraria)32

32. Thallus thin, the granules not supported by a well developed hypothallus, P+ orange-red (fumarprotocetraric acid present); rare, usually associated with Taxodium Lepraria friabilis Lendemer, K. Knudsen & Elix

32. Thallus thick, the granules supported on a well developed hypothallus, P+ orange33

33. Thallus K+ instantly intense yellow (thamnolic acid present)Lepraria aurescens Orange & Wolesley

33. Thallus K- or K+ weak, dirty yellow or brownish (other substances present)34

34. Gray to blue-green in color, never with a distinctly yellowish hue; stictic acid present; very common [if lacking TLC use this name] Lepraria finkii (B. de Lesd.) R.C. Harris

34. Blue-green to whitish in color, but always with a distinct yellowish hue; dibenzofurans present; very rare [if lacking TLC do not use this name] Lepraria vouauxii (Hue) R.C. Harris

31. Thallus continuous or areolate, not leprose, with soralia or pustules35

35. Thallus K+ intense yellow, P+ orange (thamnolic acid present)36

36. Thallus pustulose, without discrete soralia Variolaria pustulata (Brodo & W. L. Culb.) Lendemer et al.

36. Thallus not pustulose, with discrete “soralia” Variolaria trachythallina (Erichsen) Lendemer et al.

35. Thallus K- or K+ weak dirty yellow, P+ orange-red (fumarprotocetraric acid or pannarin present)37

37. Thallus pustulose, without discrete soralia, K-, P+ orange-red (pannarin present) Megalospora porphyritis (Tuck.) R.C. Harris

37. Thallus soraliate, without pustules, K- dirty yellow, P+ orange-red (fumarprotocetraric acid or succinoprotocetraric acid present) Variolaria multipunctoides (Dibben) Lendemer et al.

29. Thallus P- (above substances absent; note that occasionally atranorin gives a P+ yellow reaction when in high concentration)38

38. Thallus UV+ blue-white (2-0-methylperlatolic, alectoronic, divaricatic, perlatolic acid present)39

39. Thallus distinctly isidiate; isidia robust, tall, coralloid; alectoronic acid present Protoparmelia isidiata Diederich, Aptroot & Sérus.

39. Thallus sorediate, pustulose, leprose or with fragile isidioid soredia that are never tall and coralloid40

40. Thallus leprose, forming extensive continuous colonies; soralia absent; divaricatic acid present Lepraria hodkinsoniana Lendemer

40. Thallus continuous or areolate, not forming extensive continuous colonies; soralia, pustulose soralia or isidioid soredia present41

41. Thallus K+ yellow (atranorin present), sphaerophorin present; rare in the MACP Haematomma americanum Staiger & Kalb

41. Thallus K- (atranorin absent); 2-0-methylperlatolic acid or perlatolic acid present; common in the MACP42

42. Thallus green to green brown, forming small rosettes; discrete soralia present; perlatolic acid present; uncommon Ropalospora viridis (Tønsberg) Tønsberg

42. Thallus creamy white to blue-gray with a yellowish cast; pustulose soralia or isidioid soredia present; 2-0-methylperlatolic acid present; commonLoxospora confusa Lendemer

38. Thallus UV-43

43. Thallus sorediate or leprose (composed of granules)44

44. Thallus scruffy and dirty green in appearance, composed of goniocysts; photobiont micareoid, cells <7 μm in diameter45

45. Micareic acid present (TLC required!) Micarea prasina Fr.

45. Methoxymicareic acid present (TLC required!) Micarea micrococca (Körb.) Gams ex Coppins

44. Thallus not as above, with soredia or granules; photobiont coccoid, cells >7 μm in diameter46

46. Thallus leprose, composed entirely of granules and never with a shiny prothallus Lepraria harrisiana Lendemer

46. Thallus areolate or continuous, soraliate47

47. Thallus K-48

48. Soralia KC-, not bitter tasting (also C-, P-, and UV-; lacking secondary compounds) Variolaria ophthalmiza (Nyl.) Darb.

48. Soralia KC+ fleeting purple, bitter tasting (picrolichenic acid present) Variolaria amara Ach.

47. Thallus K+ yellow (atranorin present)49

49. Zeorin present (TLC required!)50

50. Thallus immersed, with soralia eroding from the substrate Lecanora nothocaesiella R.C. Harris & Lendemer

50. Thallus superficial, with coarse pustular soraliasorediate morph of Brigantiaea leucoxantha

49. Zeorin absent (TLC required!)51

51. Caperatic acid present52

52. Placodiolic acid group substances present; thallus usually with a fibrous white prothallus Haematomma guyanense Staiger & Kalb

52. Placodiolic acid absent; thallus without a fibrous white prothallusLecanora sp.

51. Caperatic acid absent53

53. Thallus continuous, creamy white, with discrete soralia; placodiolic acid group substances present Haematomma guyanense Staiger & Kalb

53. Thallus areolate, green, with diffuse to irregular soralia; placodiolic acid absent sterile sorediate crust sp.

43. Thallus isidiate, blastidiate, or composed of tiny areoles; soredia and granules absent54

54. On old wood, sand or organic matter55

55. Thallus composed of tiny, convex, light brown areoles [Placynthiella oligotropha (J.R. Laundon) Coppins & P. James]

55. Thallus more-or-less a continuous film of minute dark-brown to blackish areoles[Placynthiella uliginosa (Schrad.) Coppins & P. James]

54. On bark56

56. Thallus without a distinct white prothallus, areolate with the areoles developing minute globose to +/- flattened blastidiaRinodina papillata H. Magn.

56. Thallus with a distinct white prothallus, areolate with the areoles developing lobules, proliferations or isidia57

57. Areoles developing lobules or flattened, overlapping marginal proliferations58

58. Areoles large, squamulose and resembling a foliose lichen, developing lobules Phyllopsora parvifolia (Pers.) Müll. Arg.

58. Areoles small, not squamulose or resembling a foliose lichen, developing minute marginal overlapping proliferations Phyllopsora confusa Swinsc. & Krog

57. Areoles developing coralloid isidia or short globose isidia59

59. Isidia short, globose; thallus usually pale greenish gray Phyllopsora confusa Swinsc. & Krog

59. Isidia tall, cylindrical to coralloid; thallus distinctly green without a hint of gray [Phyllopsora corallina (Eschw.) Müll. Arg.]


1. Ascospores hyaline2

2. Ascospores submuriform or muriform3

3. Perithecia pale white; thallus scurfy, greenish, composed of goniocysts Psoroglaena dictyospora (Orange) H. Harada

3. Perithecia tan or black; thallus thin and indistinct, not as above4

4. Perithecia tan, inconspicuous; ostiole apical; ascospores 16–22 × 9–12; thallus concolorous with the substrate, UV-Topelia aperiens P. M. Jørg. & Vězda

4. Perithecia black, conspicuous; ostiole lateral; ascospores 40–55 × 15–20 μm; thallus gray to white, ecorticate, often patchily UV+ bright yellow (lichexanthone often present) Polymeridium proponens (Nyl.) R.C. Harris

2. Ascospores transversely septate5

5. Ascospores 4 or more celled6

6. Ascospores 4-celled7

7. Photobiont present8

8. Perithecia conspicuous, solitary, naked, and strongly raised above the thallus Trypethelium tropicum (Ach.) Müll. Arg.

8. Perithecia not conspicuously solitary and raised above the thallus surface9

9. Perithecia aggregated in raised pseudostroma; pseudostromata brown and with yellow pigmented tissue in the space between the peritheciaBathelium carolinianum (Tuck.) R.C. Harris

9. Perithecia not aggregated into raised pseudostroma10

10. Perithecia blackPolymeridium subcinereum (Nyl.) R.C. Harris

10. Perithecia light red-brown to flesh colored11

11. Perithecial wall distinctly red in section; ascospores 8 per ascusSegestria leptalea (Durieu & Mont.) R.C. Harris

11. Perithecial wall light brown in section; ascospores many per ascus Thelopsis rubella Nyl.

7. Photobiont absent; rarely 4-celled12

12. Ascospores 17–20 × 5–7 μmMycoporum eschweileri (Müll. Arg.) R.C. Harris

12. Ascospores 20–27 × 6.5–8 μmMycoporum lacteum (Ach.) R.C. Harris

6. Ascospores 6 or more celled13

13. Perithecia aggregated in conspicuous pseudostromata that often become somewhat raised; thallus bronze or light brown in colorTrypethelium virens Tuck.

13. Perithecia solitary, not aggregated in conspicuous pseudostroma14

14. Perithecia covered with abundant black apical setae; thallus with minute isidia sparsely presentTrichothelium americanum Lendemer

14. Perithecia without black apical setae; thallus not isidiate15

15. Ascospores 80–150 μmm long16

16. Ascospores narrow, 3–5 μm wide, filiform; perithecial wall dark purple-black in section Pseudosagedia rhaphidosperma (Müll. Arg.) R.C. Harris

16. Ascospores broad, 10–15 μm wide, clavate; perithecial wall brown to orange in sectionPorina heterospora (Fink) R.C. Harris

15. Ascospores <70 μm long17

17. Thallus white; ascospores fusiform to ellipsoid, 6-8 celled; rare Polymeridium quinqueseptatum (Nyl.) R.C. Harris

17. Thallus gray to brown; ascospores clavate, 8-13 celled; common Pseudosagedia cestrensis (Tuck.) R.C. Harris

5. Ascospores 2-celled18

18. Ascospores 32 per ascus, 6.5–9 × 2–3 μm; parasitic on algal colonies; rarely collected Epigloea pleiospora Döbbeler

18. Ascospores 8 per ascus, size variable; not parasitic on algal colonies; more common19

19. Perithecia arranged in minute compound aggregations that are often flattened; photobiont absent20

20. Ascospores 17–20 × 5–7 μmMycoporum eschweileri (Müll. Arg.) R.C. Harris

20. Ascospores 20–27 × 6.5–8 μmMycoporum lacteum (Ach.) R.C. Harris

19. Perithecia solitary, not arranged in minute aggregations; photobiont present except in Arthopyrenia21

21. Ascospores distinctly uniseriate within the ascus, with a median septum22

22. Thallus UV+ yellow (lichexanthone present) Anisomeridium biformoides R.C. Harris

22. Thallus UV- (lichexanthone absent)23

23. Ascospores 18–27 × 9–13 μmAcrocordia gemmata (Ach.) A. Massal.

23. Ascospores 10–18 × 4–7 μm Anisomeridium biforme (Borrer) R.C. Harris

21. Ascospores biseriate or irregularly arranged, with a submedian or median septum24

24. Ascospores 20–30 × 7–10 μm; photobiont sparsely present25

25. On hardwoods; ascomata superficial Arthopyrenia cinchonae (Ach.) Müll. Arg.

25. On Taxodium; ascomata immersedArthopyrenia taxodii R.C. Harris

24. Ascospores <20 μm long; photobiont distinctly visible26

26. Ascospores with submedian septum; paraphyses branching and anastomosing27

27. Ascospores narrowly ellipsoid, 14–20 × 4–6 μm; common Anisomeridium polypori (Ellis & Everh.) M. E. Barr

27. Ascospores broadly ellipsoid, 14–18 × 6–8 μm; rare Anisomeridium anisolobum (Müll. Arg.) Aptroot

26. Ascospores with a median septum; paraphyses not branching and anastomosing28

28. Ascospores 15–25 × 4–6 μmStrigula americana R.C. Harris

28. Ascospores 12–17 × 4–7 μm Strigula viridiseda (Nyl.) R.C. Harris

1. Ascospores brown29

29. Ascospores muriform30

30. Ascospores 130–225 × 38–50 μm long, 2 per ascus Anthracothecium nanum (Zahlbr.) R.C. Harris

30. Ascospores <70 μm long, 8 per ascus31

31. Ascospores small, 30–42–[53] × 11–15 μm; known from nearby Great Dismal Swamp [Pyrenula thelomorpha Tuck.]

31. Ascospores larger, 45–70 × 16–30 μm32

32. Perithecia aggregated into groups; ostiole lateral or apical; rare Pyrenula ravenelii (Tuck.) R.C. Harris

32. Perithecia solitary; ostiole apical; commonPyrenula leucostoma Ach.

29. Ascospores transversely septate33

33. Perithecia aggregated and fused into black pseudostromaPyrenula anomala (Ach.) R.C. Harris

33. Perithecia solitary, not fused into black pseudostroma34

34. Thallus ecorticate, white; perithecia small, black; gestalt resembling an Anisomeridium Pyrenula microcarpa Müll. Arg.

34. Thallus corticate, brown or greenish brown; perithecia not as above35

35. Ascospores >25 μm long36

36. Thallus usually red pigmented, at least near the perithecia; common Pyrenula cruenta (Mont.) Vain.

36. Thallus not red pigmented; rarePyrenula punctella (Nyl.) Trevis.

35. Ascospores <25 μm long37

37. Ascospores with the terminal lumina directly against the exospores, often citriform (resembling a lemon); perithecia not very large38

38. Thallus UV+ yellow (lichexanthone present); often on bases and boles of treesPyrenula pseudobufonia (Rehm) R.C. Harris

38. Thallus UV- (lichexanthone absent); often on the stems of shrubs Pyrenula citriformis R.C. Harris

37. Ascospores with the terminal lumina not directly against the exospore, not citriform (not resembling a lemon)39

39. Ostiole lateral; thallus thick, mostly covering the perithecia Pyrenula microtheca R.C. Harris

39. Ostiole apical; thallus thin or thick40

40. Perithecia small, 0.3–0.4 mm in diameter; thallus thin Pyrenula aspistea Ach.

40. Perithecia very large, 0.7–1.2 mm in diameter; thallus thick and shiny41

41. Hymenium densely inspersed with oil droplets Pyrenula mamillana (Ach.) Trevis.

41. Hymenium not inspersedPyrenula santensis (Nyl.) Müll. Arg.


1. Apothecia irregular (e.g., arthonioid) or elongate (lirelliform) in outline2

2. Epihymenium or exciple red or orange pigmented, pigments K+ purple or K+ green3

3. Pigment red, K+ green (isohypocrellin present); restricted to maritime forests Thalloloma cf. cinnabarinum (Fée) Staiger [collections often without ascospores]

3. Pigment red or orange, K+ purple4

4. Apothecia black, narrowly elongate and lirelliform; exciple carbonized laterally; orange pigment in the epihymenium, pigment K+ purpleGraphis inversa R.C. Harris

4. Apothecia red, irregularly shaped but not narrowly elongate and lirelliform5

5. Apothecia irregular in shape, with a distinct margin and disc; ascospores clavate, 4-celled, macrocephalicArthonia cinnabarina (DC.) Wallr.

5. Apothecia resembling a heap of mealy red granules; ascospores broadly ellipsoid, 6-celled, not macrocephalic6

6. Ascospores presentConiarthonia pyrrhula (Nyl.) Grube

6. Ascospores absentsterile material of Coniarthonia cf. pyrrhula (Nyl.) Grube

2. Epihymenium or exciple not red or orange pigmented, without pigments that are K+ purple or K+ green7

7. Apothecia blotchy and fleck-like, irregular in shape but not branching and lirelliform8

8. Ascospores transversely septate; photobiont coccoidArthonia albovirescens Nyl.

8. Ascospores submuriform to muriform; photobiont Trentepohlia or coccoid9

9. Ascospores submuriform, clavate, often +/- bentArthonia interveniens Nyl.

9. Ascospores muriform, ellipsoid, never bent10

10. Photobiont coccoid; ascospores remaining hyaline; common Arthonia susa R.C. Harris & Lendemer

10. Photobiont Trentepohlia; ascospores often turning slightly brownish with age; rare Arthonia ruana A. Massal.

7. Apothecia lirelliform, branching or not, but not blotchy and fleck-like11

11. Photobiont absentArthonia quintaria Nyl.

11. Photobiont present, Trentepohlia or coccoid12

12. Exciple carbonized, at least at the apex13

13. Disc with a thick, dense white pruina, pruina C+ red (lecanoric acid present) Dyplolabia afzelii (Ach.) A. Massal.

13. Disc variably pruinose, but pruina not thick and dense, C- (lecanoric acid absent)14

14. Lirellae forming dense, carbonized aggregations (pseudostroma) Glyphis cicatricosa Ach.

14. Lirellae solitary, not forming dense, carbonized aggregations (pseudostroma)15

15. Lirellae fissurine, visible as a narrow crack in the thallus; the excipular lips indistinct and poorly developed; exciple weakly carbonized and only at the apex; very rareFissurina subnitidula (Tuck.) Staiger

15. Lirellae not fissurine, with obvious excipular lips that are distinct and well developed; common16

16. Exciple completely carbonized17

17. Hymenium inspersed with oil droplets Graphis desquamescens Fée

17. Hymenium not inspersed18

18. Ascospores 12-14 celled; common Opegrapha viridis (Ach.) Behlen & Desberger

18. Ascospores 6-celled; infrequent19

19. Ascospores narrow, 3–4 μm wide; disc not exposedOpegrapha vulgata Ach.

19. Ascospores broad, 5–6 μm wide; disc exposed, often yellow-green pruinose Opegrapha varia Pers.

16. Exciple apically or laterally carbonized (i.e., not carbonized under the hypothecium)20

20. Exciple carbonized laterally, with the carbonization extending to the base of the hymenium21

21. Exciple smooth, entire, not becoming striate22

22. Hymenium inspersed with oil droplets23

23. Norstictic acid absent (TLC often needed for confirmation)24

24. Ascospores 20–40 μm long Graphis lineola Ach.

24. Ascospores 40–80 μm long Graphis intermedians Vain.

23. Norstictic acid present (TLC often needed for confirmation)25

25. Disc hidden by the closed lips Graphis cincta (Pers.) Aptroot

25. Disc exposed26

26. Disc white pruinose Graphis crebra Vain.

26. Disc not pruinose Graphis handelii Zahlbr.

22. Hymenium not inspersed27

27. Disc exposed, white pruinose Graphis scripta (L.) Ach.

27. Disc hidden by the closed lips28

28. Thallus corticate; lips epruinose Graphis pinicola Zahlbr.

28. Thallus at least partially ecorticate; lips lightly white pruinose Graphis furcata Fée

21. Exciple ridged, becoming striate29

29. Ascospores 30–65 × 7–12 μm Graphis striatula (Ach.) Spreng.

29. Ascospores 15–45 × 6–9 μm30

30. Lirellae long, abundantly branched Graphis duplicata Ach.

30. Lirellae short, little branched Graphis tenella Ach.

20. Exciple carbonized only at the apex, with the carbonization never extending more than one-third of the way to the base of the hymenium31

31. Exciple ridged, striateGraphis endoxantha Nyl.

31. Exciple smooth, entire, not striate Graphis vittata Müll. Arg.

12. Exciple not carbonized32

32. Thallus (test the lips) P+ yellow or orange (norstictic, psoromic or stictic acid present)33

33. Thallus P+ orange (concentrated on the lips; stictic acid present); very rare Acanthothecis leucoxanthoides Lendemer

33. Thallus P+ yellow (norstictic or psoromic acid present); common34

34. Thallus K+ yellow turning red, producing norstictic acid crystals (norstictic acid present); paraphyses ornamented with apical spines; rareAcanthothecis mosquitensis (Tuck.) E.A. Tripp & Lendemer

34. Thallus K- (psoromic acid present); paraphyses not ornamented with apical spines35

35. Apothecia fissurine, resembling mealy cracks of the thallus; ascospores muriform; throughout the DRBH Fissurina columbina (Tuck.) Staiger

35. Apothecia lecanorine, resembling irregularly shaped Lecanora; ascospores transversely septate; coastal maritime forests Enterographa anguinella (Nyl.) Redinger

32. Thallus P-36

36. Apothecia and thallus C+ pink in section (gyrophoric acid present) Arthonia anglica Coppins

36. Apothecia and thallus C- in section (gyrophoric acid absent)37

37. Thallus UV+ bright yellow, though often patchily so (lichexanthone present)Thalloloma hypoleptum (Nyl.) Staiger

37. Thallus UV- (lichexanthone absent)38

38. Lirellae fissurine, resembling a crack in the substrate39

39. Thallus blue-green to green-brown, thick and shiny40

40. Lirellae without distinct swollen lips; rare Fissurina incrustans Fée

40. Lirellae with distinct swollen lips41

41. Lirellae more-or-less immersed in the substrate; ascospores 4-celled; common Fissurina insidiosa C. Knight & Mitt.

41. Lirellae more-or-less sessile; ascospores submuriform; rare Fissurina scolecitis (Tuck.) Lendemer

39. Thallus white or gray to blue-green, thin and often dull42

42. Ascospores submuriform to muriform43

43. Ascospores large, 1 per ascus, I-, 80–100 μm long; thallus white, often flaky and appearing sorediate or schizidiate Fissurina cypressi (Müll. Arg.) Lendemer

43. Ascospores small, 6-8 per ascus, I+ violet, 15–30 μm long; thallus blue-gray to white, not flaky and appearing to have lichenized diaspores Fissurina alligatorensis Lendemer & R.C. Harris

42. Ascospores transversely septate44

44 Lirellae with white, crumbling margins; paraphyses with apical ornamentation Acanthothecis paucispora Lendemer & R.C. Harris

44. Lirellae slit-like and not with crumbling margins; paraphyses not ornamented at the tips45

45. Exciple apically carbonized, the carbonization visible as a dark area near the crack exposing the disc; rare Fissurina subnitidula (Tuck.) Staiger

45. Exciple not carbonized; common Fissurina illiterata (R.C. Harris) Lendemer

38. Lirellae not as above46

46. Ascospores muriform; photobiont coccoid Arthonia susa R.C. Harris & Lendemer

46. Ascospores transversely septate; photobiont Trentepohlia or coccoid47

47. Ascospores 6-celled, with the end two cells enlarged and the middle four cells distinctly narrowed; photobiont Trentepohlia Arthonia rubella (Fée) Nyl.

47. Ascospores 8-12-celled, cells more-or-less equal in size; photobiont coccoidArthonia albovirescens Nyl.

1. Apothecia circular in outline, regular in shape, neither irregular nor elongate48

48. Ascospores muriform49

49. Apothecia bright orange, discoid, not opening through a pore Brigantiaea leucoxantha (Spreng.) R. Sant. & Hafellner

49. Apothecia gray or brown, thelotremoid, opening through a pore50

50. Exciple carbonized, carbonized columella often present51

51. Thallus P+ yellow (subpsoromic acid present); rare in DRBH Ocellularia praestans (Müll. Arg.) Hale

51. Thallus P- (subpsoromic acid absent); common in DRBH Ocellularia sanfordiana (Zahlbr.) Hale

50. Exciple not carbonized, carbonized columella absent52

52. Ascospores 1 per ascus, >100 μm long, often turning brownish at maturity Thelotrema monospermum R.C. Harris

52. Ascospores 6-8 per ascus, 20–90 μm long, not turning brown at maturity53

53. Ascospores small, obtuse ellipsoid, 20–40 μm long; apothecia tiny, resembling a StictisThelotrema defectum R.C. Harris

53. Ascospores large, ellipsoid, 60–90 μm long; apothecia large, not resembling a StictisThelotrema adjectum Nyl.

48. Ascospores simple or transversely septate54

54. Ascospores simple55

55. Asci polysporous; ascospores many per ascus (>100), small globose or ellipsoid56

56. Thallus yellow, K+ purplePiccolia nannaria (Tuck.) Lendemer & Beeching

56. Thallus indistinct or gray-brown, K-57

57. Apothecia immersed, not lecanorine, resembling a white crater and appearing to open through a pore, K-Ramonia microspora Vězda

57. Apothecia sessile not immersed in the substrate58

58. Apothecia lecanorine, with a distinct thalline margin, medulla of the margin KC+ purple; commonMaronea polyphaea H. Magn.

58. Apothecia biatorine, without a thalline margin, margin KC-; rare Albemarlea pamlicoensis Lendemer & R.C. Harris

55. Asci not polysporous, ascospores 8 per ascus or fewer, size and shape various59

59. Apothecia contained in dome-shaped warts, tipped with an ostiole (Pertusaria)60

60. Medulla K+ yellow turning red, producing norstictic acid crystals (norstictic acid present)61

61. Ascospores 8 per ascusPertusaria propinqua Müll. Arg.

61. Ascospores 2 per ascusPertusaria neoscotica I.M. Lamb

60. Medulla K- or K+ yellow, but not K+ yellow turning red (norstictic acid absent)62

62. Thallus UV+ bright yellow (lichexanthone present) Pertusaria paratuberculifera Dibben

62. Thallus UV+ orange, UV+ dull pink, or UV-63

63. Ascospores 8 per ascus; ostiolar area usually yellow64

64. Medulla P+ orange (stictic acid present); ostiole typically raised, resembling a nipplePertusaria texana Müll. Arg.

64. Medulla P- (variolaric acid present); ostiole typically depressed, not resembling a nipplePertusaria epixantha R.C. Harris

63. Ascospores 2-4 per ascus; ostiolar area usually not yellow65

65. Thallus UV+ bright orange (thiophanic acid present) Pertusaria pustulata (Ach.) Duby

65. Thallus UV- or UV+ dull pink or orange (thiophaninic acid absent)66

66. Medulla P+ orange or red (succinoprotocetraric or stictic acid present)67

67. Ascospores 2 per ascus; warts compound, flattened; succinoprotocetraric acid present Pertusaria subpertusa Brodo

67. Ascospores (2-)4 per ascus; warts simple to compound, hemispherical; stictic acid present Pertusaria tetrathalamia (Fée) Nyl.

66. Medulla P- (succinoprotocetraric or stictic acid absent)68

68. Hymenium densely inspersed with oil droplets; warts usually sessilePertusaria sinusmexicani Dibben

68. Hymenium not inspersed; warts usually immersed Pertusaria obruta R.C. Harris

59. Apothecia not contained in dome-shaped warts, not tipped with an ostiole69

69. Apothecia lecanorine, with a distinct thalline margin (note that Variolaria species with usually sterile, convex, densely pruinose apothecia that resemble soralia also key out here)70

70. Margin and disc of apothecia C+ pink-redOchrolechia africana Vain.

70. Margin and disc of apothecia C- or C+ yellow-orange71

71. Epihymenium C+ yellow-orange (xanthones present)72

72. Disc densely white pruinose; apothecia K+ yellow turning red (norstictic acid) in sectionLecanora subpallens Zahlbr.

72. Disc lightly to densely orange-yellow pruinose; apothecia K+ yellowish (atranorin) in sectionLecanora louisianae B. de Lesd.

71. Epihymenium C- (xanthones absent)73

73. Discs densely white pruinose74

74. Ascospores 8 per ascus, small, <20 μm long Lecanora caesiorubella Ach. subsp. glaucomodes

74. Ascospores 1-2 per ascus, large, >50 μm long75

75. Pruina P+ orange or red (haemathamnolic, succinoprotocetraric, and thamnolic acid present)76

76. Thallus UV-; pruina K- or K+ dirty yellowish-brownVariolaria multipunctoides (Dibben) Lendemer et al.

76. Thallus UV+ bright yellow (lichexanthone present); pruina K+ intense yellow turning brown or reddish-brown77

77. Pruina K+ intense yellow almost instantly turning a dark reddish-brown; common Variolaria commutata (Mull. Arg.) ined.

77. Pruina K+ intense yellow slowly turning dirty brown; rare Variolaria trachythallina (Erichsen) Lendemer et al.

75. Pruina P- (other substances present)78

78. Pruina K+ lavender (hypothamnolic acid present) Variolaria hypothamnolica (Dibben) ined.

78. Pruina K- (hypothamnolic acid absent)79

79. Pruina KC+ violet (picrolichenic acid present)Variolaria amara Ach.

79. Pruina KC- (picrolichenic acid absent) Variolaria ophthalmiza (Nyl.) Darb.

73. Discs not epruinose or sparsely pruinose80

80. Thallus green; apothecial disc greenish-yellow to intense yellow (usnic acid present)81

81. Apothecial margin ecorticate, granular Lecanora strobilina (Spreng.) Kieff.

81. Apothecial margin corticate, smooth Lecanora cupressi Tuck.

80. Thallus gray; apothecial disc reddish-brown to purple-brown (usnic acid absent)82

82. Apothecial margins ecorticate, stark white contrasting against the discLecanora imshaugii Brodo

82. Apothecial margins corticate, not as above83

83. Disc dark purple-black; hymenium purple; medulla UV+ blue-white (alectoronic acid present) Tephromela atra (Huds.) Hafellner

83. Disc reddish-brown; hymenium hyaline; medulla UV- (alectoronic acid absent)84

84. Epihymenium P+ orange-red (pannarin present); infrequent in inland swamps Lecanora cinereofusca H. Magn.

84. Epihymenium P- (pannarin absent); common throughout DRBH85

85. Disc epruinose; epihymenium with fine POL+ crystals; very common Lecanora hybocarpa (Tuck.) Brodo (when in doubt, choose this one)

85. Disc weakly white pruinose; epihymenium with coarse POL+ crystals; rareLecanora chlarotera Nyl.

69. Apothecia not lecanorine, without a distinct thalline margin86

86. Apothecia bright red, K+ intense red-purple (russulone present) Ramboldia russula (Ach.) Kalb, Lumbsch & Elix

86. Apothecia tan to brown, never bright red and K+ intense purple (russulone absent)87

87. Thallus KC+ yellow-orange, UV+ orange (xanthones present)88

88. Thallus granular, leprose; apothecia dark, blackish, epruinose Pyrrhospora sp.

88. Thallus continuous, smooth to verruculose; apothecia reddish to tan, highly variable in color, frequently pruinose Pyrrhospora varians (Ach.) R.C. Harris

87. Thallus KC-, UV- (xanthones absent)89

89. Thallus scurfy, green, granular, composed of goniocysts; apothecia pallid or tan90

90. Micareic acid present [TLC required]Micarea prasina Fr.

90. Methoxymicareic acid present [TLC required] Micarea micrococca (Körb.) Gams ex Coppins

89. Thallus thin and indistinct, continuous or areolate, but not scurfy green and granular91

91. Thallus not evident; apothecia brownish-red; sporodochia absent; restricted to conifer wood in maritime forests Agyrium rufum (Pers.) Fr.

91. Thallus evident, areolate or continuous; apothecia dark brown-black; sporodochia usually present; on old large logs of conifers in swampy habitats and maritime forests Xyleborus nigricans R.C. Harris & Lendemer

54. Ascospores transversely septate92

92. Apothecial disc red or yellow-orange pigmented, pigment K+ purple, pink or green93

93. Thallus on concrete; apothecia yellow orangeCaloplaca feracissima H. Magn.

93. Thallus on bark or wood94

94. Apothecia orange; ascospores polarilocular, with thickened walls and angular luminaCaloplaca flavorubescens (With.) J.R. Laundon

94. Apothecia red; ascospores multi-celled with regular septa, not polarilocular95

95. Apothecia not lecanorine, erumpent, K+ green, often with remains of small bark flaps near the margins; rare and restricted to maritime forests Thalloloma cf. cinnabarinum (Fée) Staiger

95. Apothecia lecanorine, not erumpent, K+ red-pink or purple, always without bark flaps; common throughout the DRBH96

96. Epihymenium K+ red-pink (russulone present); medulla UV+ blue-white (sphaerophorin present); restricted to maritime forests Haematomma persoonii (Fée) A. Massal.

96. Epihymenium K+ purple (haematommone present); medulla UV- (sphaerophorin absent); primary in inland swamps97

97. Placodiolic acid present (TLC required); common Haematomma accolens (Stirt.) Hillmann

97. Pseudoplacodiolic acid and isopseudoplacodiolic acid present (TLC required); rareHaematomma flexuosum Hillmann

92. Apothecial disc not red or yellow-orange pigmented or the pigment, if present, not K+ purple or pink98

98. Apothecia large, erumpent, with large ragged bark flaps along the margins; disc pruinose Thelotrema dilatatum (Müll. Arg.) Hale

98. Apothecia not large and erumpent with large ragged bark flaps99

99. Apothecia thelotremoid, and opening through a distinct pore100

100. Ascospores large, >100 μm long; medulla C+ pink Ocellularia americana Hale

100. Ascospores small, <50 μm long; medulla C-101

101. Thallus ecorticate, poorly developed; common Thelotrema subtile Tuck. (when in doubt, choose this one)

101. Thallus corticate, well-developed; rareThelotrema lathraeum Tuck.

99. Apothecia discoid, not opening through a pore102

102. Ascospores 2-celled103

103. Photobiont Trentepohlia; apothecia yellowish-orange Coenogonium luteum (Dicks.) Kalb & Lücking

103. Photobiont coccoid; apothecia reddish brown or pale to tan104

104. Ascospores polarilocular, with thickened walls and angular luminaCaloplaca camptidia (Tuck.) Zahlbr.

104. Ascospores not polarilocular, with regular septa105

105. Apothecia distinctly bicolored, with a white margin contrasting against the dark blue-black disc; rare Megalaria alligatorensis Lendemer & R.C. Harris

105. Apothecia not bicolored, the margin (if present), concolorous with the disc; common106

106. Apothecia reddish-brown; thallus smooth or thin areolate Catinaria atropurpurea (Schaer.) Vězda & Poelt

106. Apothecia tan or pallid; thallus scurfy, green, granular, composed of goniocysts107

107. Apothecia raised, strongly stipitate, with a roughened surface due to the protrusion of paraphyses and asci from the hymenium; infrequent Vezdaea leprosa (P. James) Vězda

107. Apothecia not raised and stipitate, with a smooth surface; common108

108. Micareic acid present [TLC required] Micarea prasina Fr.

108. Methoxymicareic acid present [TLC required]Micarea micrococca (Körb.) Gams ex Coppins

102. Ascospores >2-celled109

109. Ascospores fragile, readily fragmenting into part spores110

110. Ascospores <30 μm long, 4-6 celled; on Taxodium; very rare Bactrospora brevispora R.C. Harris

110. Ascospores >30 μm long, 10-20 celled; on various substrates111

111. Ascospores 35–50 × 3 μm; common Bactrospora carolinensis (Ellis & Everh.) R.C. Harris

111. Ascospores 80–90 × 7–10 μm; infrequent Bactrospora lamprospora (Nyl.) Lendemer

109. Ascospores not fragile, typically remaining intact and not forming part spores112

112. Ascospores needle shaped, <3 μm wide113

113. Ascospores short, <20 μm long114

114. Pycnidia stipitate; thallus UV+ blue-white (lobaric acid present) and usually P+ orange-red (fumarprotocetraric acid present); common on Pinus Micarea neostipitata Coppins & P. May

114. Pycnidia not stipitate; thallus UV- and P- (lobaric acid and fumarprotocetraric acid absent); rare and restricted to Chamaecyparis and Taxodium Micarea chlorosticta (Tuck.) R.C. Harris

113. Ascospores longer, >20 μm long115

115. On concreteBacidina egenula (Nyl.) Vezda

115. On bark116

116. Hypothecium dark brown117

117. Epihymenium K+ purple; hypothecium orange-brown pigmented, pigment K- Bacidia helicospora S. Ekman

117. Epihymenium K-; hypothecium brown pigmented, pigment K- or K+ rose-red118

118. Brown pigmented portions of the exciple diffuse K+ rose-red; rare Bacidia diffracta S. Ekman

118. Brown portions of the exciple K- or K+ more intense brown-purple; common119

119. Epihymenium blue-green pigmented; hypothecium and exciple dark purple-brown pigmented; apothecia dark black in color Bacidia schweinitzii (Fr. ex Tuck.) A. Schneid.

119. Epihymenium hyaline; hypothecium and exciple reddish-brown pigmented; apothecia reddish brown in color Bacidia schweinitzii (Fr. ex Tuck.) A. Schneid. (brown color form)

116. Hypothecium pale yellowish to hyaline120

120. Epihymenium with pigment forming distinct caps over the paraphyses; primarily coastal Bacidia heterochroa (Müll. Arg.) Zahlbr.

120. Hymenium with evenly dispersed pigment, or not pigmented121

121. Apothecia dark brown; epihymenium K+ purple Bacidia helicospora S. Ekman

121. Apothecia pale, pallid; epihymenium K-Bacidina spp. not treated here

112. Ascospores fusiform, ellipsoid or otherwise, but >3 μm wide and not needle shaped122

122. Margins of apothecia fuzzy, white, byssoid123

123. Hypothecium pale; apothecial discs pale yellow, thallus UV+ dull orange (xanthones) present) Byssoloma meadii (Tuck.) S. Ekman

123. Hypothecium dark; apothecial discs dark blue-gray or dark brown, thallus UV- Byssoloma leucoblepharum (Nyl.) Vain.

122. Margins of apothecia smooth, not fuzzy and byssoid124

124. Apothecial discs yellow or greenish pruinose Cresponea flava (Vain.) Egea & Torrente

124. Apothecial discs epruinose or brown pruinose125

125. Apothecia resembling tiny black urns; discs brown pruinose Glyphis scyphulifera (Ach.) Staiger

125. Apothecia not resembling tiny black urns; discs epruinose126

126. Apothecial discs white, C+ pink in section (gyrophoric acid present); restricted to inland swampsMicarea peliocarpa (Anzi) Coppins & R. Sant.

126. Apothecial discs dark brown, C- (gyrophoric acid absent); mostly in coastal maritime forests127

127. Ascospores 4-celled; thallus thick, apothecia immersed in the thallus Mazosia carnea (Eckfeldt) Aptroot & M. Cáceres

127. Ascospores 6-celled; thallus thin, apothecial sessile Schismatomma cf. rappii (Zahlbr.) R.C. Harris


1. Apothecia elongate in outline, lirelliform2

2. Ascospores transversely septate, 4-10 celled3

3. Ascospores 6-10 celled4

4. Lirellae distinctly elongate, often weakly branched, not erumpent and surrounded by ragged bark flaps; ascospores 6-celled[Phaeographis/Leiorreuma sp.]

4. Lirellae circular to distinctly elongate, strongly erumpent and often surrounded by ragged bark flaps; ascospores 6-10 celled5

5. Coastal maritime forests; lirellae circularPhaeographis lobata (Eschw.) Müll. Arg.

5. Inland swamps, often in the canopy; lirellae elongate Phaeographis erumpens (Nyl.) Müll. Arg.

3. Ascospores 4-celled6

6. Hymenium not inspersed; medulla P+ yellow or orange (norstictic and/or stictic acid present); exciple weakly carbonized at the apex, if at all Phaeographis brasiliensis (A. Massal.) Kalb & Matthes-Leicht

6. Hymenium inspersed with oil droplets; medulla P- (norstictic/stictic acid absent); exciple variably carbonized7

7. Lirellae forming aggregations (pseudostroma) of variable branching density where the hymenium becomes cracked and divided; pseudostromata surrounded by distinctly whitish tissue that differs markedly in color and texture from the surrounding thallusSarcographa tricosa (Ach.) Müll. Arg.

7. Lirellae not forming aggregations (pseudostroma) that are surrounded by distinctly whitish tissue that differs markedly in color and texture from the surrounding thallus8

8. Exciple apically carbonized; rare in DRBH Phaeographis inusta (Ach.) Müll. Arg.

8. Exciple completely carbonized, with the carbonization often penetrating into the hypothecium; common in DRBHLeiorreuma sericeum (Eschw.) Staiger

2. Ascospores submuriform to muriform9

9. Ascospores submuriform, 25–30 × 10–12 μm; lirellae elongate, often branching, with a distinct white thalline margin, not erumpent; throughout DRBHLeiorreuma explicans (Fink) Lendemer

9. Ascospores muriform, 90–150 × 25–35 μm; lirellae circular, never branching, erumpent and surrounded by ragged bark flaps; restricted to mature maritime forests Phaeographis oricola Lendemer & R.C. Harris

1. Apothecia circular in outline.10

10. Ascospores muriform11

11. Apothecia immersed, opening through a pore, not erumpent surrounded by ragged bark flaps; ascospores 100–150 × 25–30 μm; restricted to inland swamps Thelotrema monospermum R.C. Harris

11. Apothecia immersed, but not opening through a pore, strongly erumpent and surrounded by ragged bark flaps; ascospores 90–150 × 25–35 μm; restricted mature maritime forests Phaeographis oricola Lendemer & R.C. Harris

10. Ascospores transversely septate, < 90 × 20 μm12

12. Ascospores 4-10 celled13

13. Ascospores 4-celled; hymenium not inspersed; apothecia lecideine, small; thallus composed of minute green areoles; inland swampsBuellia vernicoma (Tuck.) Tuck.

13. Ascospores 6-10 celled; hymenium inspersed with oil droplets; apothecia erumpent, large; thallus a thick brown continuous crust; maritime forests Phaeographis lobata (Eschw.) Müll. Arg.

12. Ascospores 2-celled14

14. Hypothecium hyaline15

15. Ascospores (18–)24–27(−34) × (11–)13–15(−17) μm; rare Rinodina dolichospora Malme

15. Ascospores (12–)17–18(−23) × (6–)8–9(−10) μm; common Rinodina maculans Müll. Arg.

14. Hypothecium brown16

16. Ascospores 16-32 per ascusAmandinea polyspora (Willey) E. Lay & P. May

16. Ascospores 8 or fewer per ascus17

17. Thallus K+ yellow turning red, with norstictic acid crystals in section (norstictic acid present)18

18. Ascospores 10–15 × 5–7 μm; typically on trunks at base and boles Buellia stillingiana J. Steiner

18. Ascospores 16–23 × 6–10 μm; typically on branches in the canopy or on small stems of shrubs19

19. Ascospores with thickened walls and angular lumina; on Chamaecyparis; rareBuellia imshaugiana R.C. Harris

19. Ascospores not with thickened walls and angular lumina; on diverse substrates including Chamaecyparis; common Buellia curtisii (Tuck.) Imshaug

17. Thallus K- or K+ yellowish, but not turning red with norstictic acid crystals in section (norstictic acid absent)20

20. Apothecial section with K+ pink or orange-red pigments in the epihymenium or exciple21

21. Photobiont present; thallus composed of minute green areoles; epihymenium K+ fleeting pink; exciple K-Buellia elizae (Tuck.) Tuck.

21. Photobiont absent; thallus not evident; epihymenium K-; exciple K+ strong orange-redgen et sp. nov. aff. Schrakia

20. Apothecial section without K+ pink or orange red pigments in the epihymenium or exciple22

22. Thallus composed of minute green areoles containing bright pink pigment (pigment often difficult to observe without a compound microscope), P+ yellow (baeomycesic acid present) Gassicurtia acidobaeomyceta Marbach

22. Thallus not as above, without a pigment, P-23

23. Ascospores with thickened walls and angular lumina; restricted to maritime forestsHafellia sp.

23. Ascospores without thickened walls and angular lumina; distribution various24

24. Ascospores <14 μm long, smooth, not ornamented25

25. Apothecia erumpent, retaining a thalline margin, at least when young; coastal maritime forests Amandinea milliaria (Tuck.) P. May & Sheard

25. Apothecia not erumpent, without a thalline margin; throughout Amandinea punctata (Hoffm.) Coppins & Scheid.

24. Ascospores >14 μm long, rough, ornamented27

26. Thallus barely evident, always lacking soralia; ascospores 15–18 × 10–12 μm; restricted to maritime forests Amandinea langloisii Marbach

26. Thallus evident, composed of thin green areoles, often with a few sparse yellow soralia at the margins; ascospores 14–19 × 9–12 μm; common throughout DRBHrare abundantly fertile forms of Buellia wheeleri R.C. Harris

James C. Lendemer, Richard C. Harris, and Ana Maria Ruiz "A Review of the Lichens of the Dare Regional Biodiversity Hotspot in the Mid-Atlantic Coastal Plain of North Carolina, Eastern North America," Castanea 81(1), 1-77, (1 March 2016).
Received: 22 October 2015; Accepted: 1 March 2016; Published: 1 March 2016
barrier island
biodiverse understudied groups
obligate symbionts
Outer Banks
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