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Opuscula Philolichenum 21: 33–181. 2022. [authorship corrected version, published 29 September 2022]
*pdf first effectively published online 14June2022 via (http://sweetgum.nybg.org/philolichenum/)
Getting to know our biomonitor neighbours: urban
lichens and allied fungi of Edmonton, Alberta,
Canada
DIANE L. HAUGHLAND
1
*, ALESSANDRA HOOD
2
, DARCIE THAUVETTE
3
, SYDNEY A. TONI
4
, MING CAO
5
,
JOSEPH D. BIRCH
6
, JOSHUA WASYLIW
7
, LAURA HJARTARSON
8
, MARY VILLENEUVE
9
, ARYNN STORDOCK
10
,
DAVID A. FIELDER
11
, MEGAN LEWIS
12
, DAVID EVANS
13
, DOMINIK ROYKO
14
, RASHELL BOLDUC
15
, HAYLEY
WEBSTER
16
, JEREMY D. SINGH
17
, KRISTEN A. SCHAFER
18
, SPENCER GOYETTE
19
, HANNA E. DAVIDSON
20
AND CATHERINE SHIER
21
1
DIANE L. HAUGHLAND – Alberta Biodiversity Monitoring Institute, CW 405 Biological Sciences Building, University of
Alberta, Edmonton, Alberta, T6G 2E9, Canada. & Department of Renewable Resources, Faculty of Agricultural, Life &
Environmental Sciences, 751 General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-
mail: dianeh@ualberta.ca – *author for correspondance
2
ALESSANDRA HOOD – Royal Alberta Museum, 12845 102 Avenue, Edmonton, Alberta, T5N 0M6, Canada. – e-mail:
ajhood@ualberta.ca
3
DARCIE THAUVETTE – Alberta Biodiversity Monitoring Institute, CW 405 Biological Sciences Building, University of
Alberta, Edmonton, Alberta, T6G 2E9, Canada. – e-mail: darcie.thauvette@ualberta.ca
4
SYDNEY A. TONI – Alberta Biodiversity Monitoring Institute, CW 405 Biological Sciences Building, University of Alberta,
Edmonton, Alberta, T6G 2E9, Canada. & Royal Alberta Museum, 12845 102 Avenue, Edmonton, Alberta, T5N 0M6,
Canada. – e-mail: stoni@ualberta.ca
5
MING CAO – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751 General
Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: mcao3@ualberta.ca
6
JOSEPH D. BIRCH – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: jcooper@ualberta.ca
7
JOSHUA WASYLIW – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: wasyliw@ualberta.ca
8
LAURA HJARTARSON – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: lhjartar@ualberta.ca
9
MARY VILLENEUVE – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: mvillene@ualberta.ca
10
ARYNN STORDOCK – Royal Alberta Museum, 12845 102 Avenue, Edmonton, Alberta, T5N 0M6, Canada. – e-mail:
stordock@ualberta.ca
11
DAVID A. FIELDER – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: fielderda@gmail.com
12
MEGAN LEWIS – Department of Agricultural, Food & Nutritional Sciences, Faculty of Agricultural, Life & Environmental
Sciences, 410 Agriculture/Forestry Centre, University of Alberta, Edmonton, Alberta, T6G 2P5, Canada. & Department of
Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751 General Services Building, University
of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: mclewis@ualberta.ca
13
DAVID EVANS – Innotech Alberta – Vegreville, P.O Box 4000, Hwy 16A & 75 Street, Vegreville, Alberta, T9C 1T4,
Canada. & Alberta Biodiversity Monitoring Institute, CW 405 Biological Sciences Building, University of Alberta,
Edmonton, Alberta, T6G 2E9, Canada. – e-mail: david.evans@innotechalberta.ca
14
DOMINIK ROYKO – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: royko@ualberta.ca
15
RASHELL BOLDUC – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail:
rashell.b123@gmail.com
16
HAYLEY WEBSTER – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: hwebster@ualberta.ca
17
JEREMY D. SINGH – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: jdsingh@ualberta.ca
18
KRISTEN A. SCHAFER – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: kschafer@ualberta.ca
19
SPENCER GOYETTE – Department of Biological Sciences, Faculty of Science, CW 405, Biological Sciences Building,
University of Alberta, Edmonton, Alberta, T6G 2E9, Canada. – e-mail: sgoyette@ualberta.ca
20
HANNA E. DAVIDSON – Department of Renewable Resources, Faculty of Agricultural, Life & Environmental Sciences, 751
General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada. – e-mail: garvey@ualberta.ca
21
CATHERINE SHIER – Planning and Environment Services, Urban Planning and Economy, City of Edmonton, Edmonton
Tower, 10111 - 104 Avenue NW, Edmonton, Alberta, T5J 0J4, Canada. – e-mail: catherine.shier@edmonton.ca
34
ABSTRACT. – Here we provide one of the first detailed studies of lichen and allied fungi
diversity in a continental North American city (Edmonton, Alberta, Canada), including an annotated
checklist, images of all species, dichotomous keys, and local distribution maps. Edmonton is the
northernmost city in North America with a population of over one million, and an industrial and
transportation gateway for much of northern Canada. Lichen-based biomonitoring could be a tool to track
airborne pollutants resulting from Edmonton’s growing populace and industrial activity. The first step
towards such a program is documenting the diversity and distribution of lichens in the city. To accomplish
this, we conducted a city-wide, systematic survey of 191 sites focused on epiphytes growing on deciduous
boulevard trees. We augmented that survey with surveys of rare trees, opportunistic collections from river
valley and ravine habitats, herbarium collections, phylogenetic analyses of a subset of collections, and
observations submitted to online nature-reporting applications. We present ITS sequence barcode data for
33 species, phylogenetic analyses for Candelariaceae, Endocarpon, Flavopunctelia, the Lecanora dispersa
group, Lecidella, Peltigera, Physconia, and Punctelia, and detailed descriptions of 114 species in 47 genera
and 23 families. Two species are hypothesized to be new to North America (Endocarpon aff. unifoliatum,
Lecidella albida), twelve more are new to Alberta (Amandinea dakotensis, Bacidia circumspecta,
Candelaria pacifica, Candelariella antennaria, Heterodermia japonica, Lecania naegelii, Lecanora
sambuci, Lecanora stanislai, Lecidea erythrophaea, Peltigera islandica, Phaeocalicium aff. tremulicola,
and the introduced Xanthoria parietina), and five are putative new species to science (Physcia aff.
dimidiata, Physcia aff. stellaris, Phaeocalicium sp., Phaeocalicium aff. tremulicola, Lichenaceae sp.).
Illustrations are provided for all species to aid in verification and public outreach. Species richness was
highest in foliose lichens (48), followed by crustose and calicioid lichens and allied fungi (41), with the
lowest richness in fruticose lichens (25). We did a preliminary assessment of the suitability of species for
citizen-science biomonitoring by assessing their distribution across the city, perceptibility to the public,
identification accuracy, and, for a subset, how consistently species were surveyed by trained novices.
Compared to other urban areas where lichen diversity has been studied, Edmonton is relatively species-rich
in calicioids and Peltigera. Promising bioindicators may be limited to chlorolichens, including Caloplaca
spp., Evernia mesomorpha, Flavopunctelia spp., Phaeophyscia orbicularis, Physcia adscendens, Physcia
aipolia group, Physcia aff. stellaris, Usnea spp., and Xanthomendoza fallax. Other genera that may be
responsive to pollutants such as Cladonia and Peltigera were almost exclusively restricted to river valley
and ravine ecosystems, limiting their application as bioindicators. Some species commonly used as
biomonitors elsewhere were too rare, small, poorly developed, or obscured by more common species
locally (e.g., Candelaria concolor s.l., Xanthomendoza hasseana). The low overlap with lists of
biomonitoring species from other regions of North America illustrates the necessity of grounding
monitoring in knowledge of local diversity. Future augmentation of this list should focus on enhanced
sampling of downed wood-, conifer-, and rock-dwelling lichens, particularly crustose species. The next step
in developing a biomonitoring program will require modelling species’ responses to known air quality and
climatic gradients.
KEYWORDS. – Air quality, lichenized Ascomycetes, biomonitoring, calicioids, Candelariaceae,
cyanolichens, continental climate, detection error, Endocarpon, Flavopunctelia, floristics, Lecanora
dispersa group, Lecidella, molecular phylogenetics, Peltigera, Punctelia, survey repeatability, urban
biodiversity.
INTRODUCTION
Lichens are widespread, spanning the globe from the Arctic to Antarctica (Sancho et al. 2019),
grow on many different substrates, are long-lived, and can be studied in any season (Nimis et al. 2002).
Individual species’ responses to atmospheric conditions including anthropogenic pollutants determine
overall species diversity patterns (e.g., Matos et al. 2019). These traits and others have led to a long history
of using lichens in urban environments as biological monitors (biomonitors) of air quality (Grindon 1859,
Llop et al. 2012, Nimis et al. 2002, Nylander 1866, Pungin & Dedkov 2017). While physicochemical air
35
quality monitoring stations can provide highly accurate contaminant data, they are costly and do not
provide the high spatial resolution of multiple lichen sampling sites. Air quality stations can monitor levels
of specific air quality parameters like SO2 and NO2 from point sources, but differences in lichen
communities can indicate relative deposition and the cumulative biological effect of complex atmospheric
conditions across an entire city (McCarthy et al. 2009).
In Canada, lichens have been used as biomonitors for a variety of pollutants (reviewed by
MacDonald & Coxson 2013, Thormann 2006), but lichen surveys have failed to gain widespread traction in
biomonitoring. In 1994, the federal government created the Ecological Monitoring and Assessment
Network (EMAN) and commissioned the creation of standardized protocols for community-based
ecological and biodiversity monitoring. EMAN included a lichen-based, rapid-assessment air quality
protocol, using lichens that could be identified in the field, based largely on work in the milder climes of
the Mixedwood Plains ecozone (Marshall et al. 1999, ecozones.ca) of southeastern Canada. The EMAN
protocol was trialed in Hamilton, Ontario (McCarthy et al. 2009), but shortly thereafter EMAN was
defunded. Similar studies in continental climates are limited. In Stringer and Stringer (1974), lichen
diversity was mapped and interpreted based on putative air quality gradients in Winnipeg, Manitoba.
Locally, Case (1980) examined the response of epiphytic lichens on conifers to proximity to a sour gas
(H2S) plant approximately 200 km northwest of Edmonton, Alberta. Lee & Vitt (1974) and Elsinger et al.
(2007) examined the response of epiphytic lichens on Populus trees in proximity to a zone of industrial
activity on the outskirts of Edmonton. The province of Alberta has a long history of industrial activity,
including oil and gas extraction and refinement, pulp and paper mills, and coal mining. These industries
have been associated with air pollution including H2S, NOX, fine particulate matter, polycyclic aromatic
hydrocarbons, and trace element deposition (Alberta Capital Air Shed 2017, Government of Canada 2016,
Kindierski & Bari 2015, Makar et al. 2018). Given the prevalence of these industries and their proximity to
major population centers, there is a need to better understand how air pollution may impact local species,
such as lichens, as well as human health (e.g., Rodriquez-Villamizar et al. 2017).
Alberta has benefited from the attention of resident and visiting lichenologists (e.g., Bird 1970,
1973, Björk 2013, Haughland et al. 2018, McCune & Goward 1995, Spribille 2002, Thomson & Ahti
1994), and a province-wide biodiversity monitoring program that includes lichens (the Alberta Biodiversity
Monitoring Institute, hereafter referred to as the ABMI [2020], www.abmi.ca). However, as in many
regions, little is published on lichens in urban environments. Alberta’s capital city Edmonton is an
interesting test case for biomonitoring with lichens because it has a cold, dry, temperate climate that is not
associated with many of the lichens found to be effective biomonitors elsewhere; it is bisected by a ribbon
of river valley parks, potentially elevating urban lichen diversity; and, as in many Canadian cities, there is a
long history of planting deciduous boulevard trees that could be used as a standard substrate for lichen
biomonitoring. Lichens are colorful components of many urban landscapes, adorning boulevard trees lining
sidewalks and streets. With investment in public outreach and identification training, lichens can become
more perceptible to the public, who could be mobilized to document local lichens as potential air quality
and/or climate change biomonitors.
Here we aim to address Edmonton’s deficit of lichen knowledge by publishing the first annotated
lichen list for the city. We sought to document patterns of lichen diversity across the Edmonton area to 1)
inform future studies on lichens as citizen science biomonitors, and 2) disseminate documentation on lichen
diversity and identification to help non-specialists learn the broad groups of lichens and appreciate lichen
diversity. We conducted systematic surveys of boulevard trees across the city, rare tree surveys,
opportunistic surveys in river valley and ravine parklands, herbarium searches, online database searches,
and searches of lichen records in two public, online nature-reporting applications, iNaturalist (2020) and
NatureLynx (2020). We conducted molecular phylogenetic analyses to aid in the identification of a subset
of collections. We used these composite data to determine if species had some of the attributes of an
effective biomonitor. Lichens were deemed potential biomonitors if they were relatively abundant, widely
distributed, present in different habitats, perceptible to the public, and amenable to consistent, accurate
identification. We used online observations as a metric of perceptibility, i.e., which species and genera the
public noticed. We assessed accuracy through examination of vouchers, herbarium specimens, visual
verifications of online reports, as well as field audits of a portion of the systematic boulevard tree surveys.
This work assesses the potential for using lichens as citizen science biomonitors in Edmonton, as well as a
filter to guide future work modeling the responsiveness of promising lichen species to urban climate and air
quality gradients.
36
MATERIALS AND METHODS
Study Area. – Edmonton (53°34′24″N, 113°25′06″W, 620–720 m elevation, municipal area of
783 km2, Fig. 1), is the northernmost North American municipality with a population over one million, and
is an industrial and transportation gateway to Alberta’s northeastern oil and gas region. Industry in
Edmonton is concentrated in three main areas: along the eastern border, within the northwest portion of the
city, and from the central downtown core extending south to 41 Avenue SW. Edmonton previously had a
municipal airport approximately 2 km from the downtown core, which began as an airfield in 1927
(Griwkowsky 2017) and operated until 2013.
Edmonton’s current average airborne contaminant concentrations are estimated at 13–19 ppb
nitrogen oxides (NOx), 19.7–24.8 ppb ozone (O3), <1–1.7 ppb sulphur dioxide (SO2), and <1 ppb sour gas
(H2S) (Alberta Capital Airshed 2017). From 1998 to 2014, many pollutants trended downwards (Kindierski
& Bari 2015). Most nitrates and sulfates are hypothesized to be generated by local vehicle exhaust and
industrial activities, coal combustion from west of Edmonton, larger-scale regional activities such as oil
and gas extraction and production in Alberta, British Columbia, and Saskatchewan, and animal feeding
operations south of Edmonton to Montana, U.S.A. In 2017, Edmonton ranked 9th highest of 16 Canadian
cities for SO2, 3rd of 24 cities for NO2, 22nd of 24 cities for O3, and 5th of 24 cities for fine particulate matter
(PM2.5) (Alberta Capital Airshed 2017).
Ecologically, Edmonton is within the Parkland Natural Region and Central Parkland Natural
Subregion of Alberta, which can be summarized vegetatively as an aspen forest–grassland mosaic. The
Central Parkland is considered a transitional Natural Subregion between the Grasslands Natural Region to
the south and Boreal Mixedwoods Natural Subregion to the north (Natural Regions Committee 2006).
Edmonton thus shares climatic and vegetation characteristics with several natural regions. These
transitional characteristics are critical to the city’s overall biodiversity, as the resulting diverse plant
communities create distinct habitats and wildlife assemblages unique to the Parkland.
Native Parkland is an ecological system found only in North America (Natural Regions
Committee 2006). It is characterized by rich soils, a canopy of aspen and balsam poplar (Populus
tremuloides Michx. and Populus balsamifera L.), and a species-rich understory of shrubs. Typical habitats,
dictated by local geology, topography, soils, and hydrology, include forested river valley and ravine slopes,
riparian habitats, patches of mixedwood boreal forest, deciduous woodlands, lakes and smaller wetlands,
and small areas of grassland, with some remnant sand dune, peatland and shrubland habitats. Today,
Parkland habitat is scarce, with >78% of the Natural Region directly altered by anthropogenic activity
(ABMI 2018).
Edmonton can be divided into two distinct geomorphic areas: the tablelands and the river valley
and ravine system. The tablelands area is by far the larger of the two and is relatively flat, varying
approximately between 650–675 meters in elevation. Much of the area is under urban and industrial
development or cultivation but remnants of the historical Central Parkland, aspen-grassland mosaic remain
in small regions of the city. The river valley and ravine system occupy 10% of the land base and consist of
natural areas, including the North Saskatchewan River that runs through the middle of the city and its
associated ravines, as well as wetlands and forests that dot the tablelands above the valley. Edmonton’s
river valley constitutes the longest stretch of connected urban parkland in North America (City of
Edmonton 2013). Natural disturbances historically shaped the landscape, with most of the North
Saskatchewan River Valley reset to early successional Populus stands by a fire prior to 1879 (Wein 2006).
After the 1900s, stand age and biodiversity were mostly impacted by riparian logging, hillside tunnel
mining for coal, and gravel dredging for gold (Wein 2006).
Edmonton has a continental climate and receives most of its precipitation in the summer, with an
average 94 mm of rain in the wettest month of July and 25 mm of snow in the snowiest month of January,
and a total of 456 mm of precipitation annually (climate normals 1981-2010, Government of Canada 2010).
Daily average temperatures range from -10.4 ± 4.8°C (SD) in January to 17.7 ± 1°C in July. Average
relative humidity varies from 76% at 6:00 am to 54% at 3:00 pm.
Data sources. I. Field surveys. – We conducted three types of surveys: opportunistic surveys in
lichen-diverse areas (2013 onwards), systematic surveys of epiphytic lichens growing on the boles of
deciduous trees as part of a University of Alberta (U of A) class project (2019), and surveys of rare tree
species growing on the U of A campus (2021).
37
Figure 1. City of Edmonton, Alberta, Canada (53°34'24" N, 113°25'06" W, 670 m elevation), with lichen
survey locations and observations mapped. The Anthony Henday is a major ring road approximately
delimiting the area of focus. Not shown are two air quality stations surveyed that lie east of the city.
Edmonton is within the Parkland Natural Region, surrounded by the Boreal Natural Region to the north,
east, and west (inset map).
2019 Systematic surveys. – To systematically survey epiphytic lichens, we surveyed 191 sites
located across Edmonton. We defined the area of interest as a 440 km2 region contained within Edmonton’s
outer ring road, Anthony Henday Drive (Fig. 1). We chose a 1.5 km grid spacing to make our study
comparable to that of McCarthy et al. (2009) and because it generated a logistically feasible number of sites
(n=135) for our initial survey effort in February and March. An additional 49 sites were added to enhance
coverage wherever possible in industrial zones. We also conducted more intensive surveys near seven
permanent air quality monitoring stations for a total of 1,095 trees examined across 191 sites.
We generated a map for each site showing nearby potentially suitable trees using the Open
Edmonton Tree Database (City of Edmonton 2019) and ArcGIS. The database contains information on
>350,000 city-owned trees located in urban parks, on boulevards, and road right-of-ways. Each tree in the
database has a unique identification number, planting year, species, and diameter at breast height (DBH;
1.37 m). We targeted deciduous trees in the genera Fraxinus, Populus and, Ulmus with a planting year of
1990 because of a long history of planting these genera in Edmonton, ensuring that relatively mature trees
were available in many neighborhoods. Other deciduous species were surveyed if the target genera were
not available. The planting year was a placeholder for trees planted prior to or in 1990. However, when
38
trees are replaced by the city, the planting date and species are rarely updated. To address this unanticipated
source of error, we independently recorded the tree species (or genus where species could not be assessed),
circumference, as well as the latitude and longitude of each tree surveyed when we could not
unambiguously assign a tree a known tree identification number.
At each survey location, participants surveyed the five closest target trees with a minimum DBH
of 15 cm. At air quality monitoring stations, we increased survey intensity to 25 trees. We avoided trees
with potentially anomalously high exposure to traffic exhaust such as those at bus stops, next to
intersections, and along roads with more than one lane in each direction. Wherever possible we also
avoided trees where snow was piled more than 0.5 meters high around the base or that had significant bark
damage. Surveys were conducted between 25 February 2019 and 25 July 2019. The early survey period
was marked by below-average cold weather, with temperatures reaching -40°C with wind chill. We thus
minimized survey duration and avoided methods that required dexterity (such as attaching quadrats to trees
for frequency counts) and instead measured relative cover. Surveyors recorded the presence of each of 21
target species or genera (Table 3) on the north-, south-, east-, and west-facing sides of each tree, between
approximately 0.5 m to 1.5 meters off the ground (to avoid snow-covered or nutrient-enriched tree bases,
and to make the survey area more comparable between surveyors of different heights). The target taxa were
chosen by the senior author based on prior observation within Edmonton and because we hypothesized they
could be accurately determined by novice surveyors. Each species was scored as absent (0) or present with
a relative abundance of 1, 2, 3 or 4 (i.e., the number of sides the species was present on). While this
simplified differences in cover between species, reducing the time spent estimating cover allowed us to
scan a larger proportion of tree surface area and capture more species. When surveyors could not identify a
lichen, they collected representative small vouchers for later verification. Lichens that could be identified in
the field but which were not on the target list were added to the site list and surveyed as above. Initially,
only macrolichens were included in the target list, however, on the first day of the survey we added the
crustose genera Caloplaca and Candelariella due to their abundance and hypothesized detectability.
Samples of crustose lichens were taken at a subset of sites for later identification.
There were three groups of surveyors. The trained novices consisted of undergraduate, graduate,
and open-learning students (n=8) that had completed the first half of a course on the lichens of Alberta.
This group completed approximately 14 hours of laboratory study of lichens prior to the project, including
weekly testing that covered all but one of the species on the target list. The intermediate group included a
former student, a new lichen technician, the lichen course Teaching Assistant, and an experienced ABMI
field supervisor (n=4). All had received training and participated in some lichen-related fieldwork prior to
the project. The experts consisted of the senior author and a lichen technician each with more than ten years
of laboratory and field experience (n=2).
Participants received approximately 1–2 hours of field training on survey methodology on their
first day of monitoring by one of the experts. Experts provided additional field assistance and quality
checks. To estimate observer effects, 50 sites were audited by pairs of intermediate and expert surveyors.
Thirty-four additional sites were revisited to better document the crustose lichen flora once it was
determined the original surveyors had not captured those species (based on audits and information from
surveyors).
2021 Rare tree surveys. – Two authors (K. Schafer and J. Singh) surveyed tree species that are
relatively rare in Edmonton as part of the 2021 U of A Lichens of Alberta class. Trees growing on the U of
A North Campus were chosen because they were not represented in existing surveys, and included species
of conifers in the genera Larix, Picea, Pinus, Taxus, and Thuja, as well as deciduous species in the genera
Acer, Aesculus, Betula, Fraxinus, Quercus, Salix, Syringa, and Ulmus. In total, 44 individual trees were
surveyed, and all accessible surfaces of those trees (bole, branches, twigs) were examined for crustose,
calicioid, and macrolichens. Representative samples were taken from each tree and identified or confirmed
in the laboratory with the help of the senior author. These data were added to the combined dataset for
mapping and inclusion in the annotated species list.
Data sources. II. Historical and herbarium records. – To establish a list of lichens previously
recorded in Edmonton, we searched the literature, databases of the Royal Alberta Museum (PMAE), a
small portion of the University of Alberta (ALTA) lichen records (most of the lichen records are not
databased at this time and the herbarium was closed to visitors due to the COVID pandemic), and the
Canadian Forestry Centre in Edmonton (provided by G. Pohl, pers. comm.), as well as the Government of
39
Alberta’s online species tracking dataset BIOTICS (Government of Alberta 2020). Any specimens that
could be located were verified or reidentified. We did not consider a 1:1 taxonomic change a
misidentification. For example, a correctly determined Melanelia albertana or Parmelia albertana
(currently Melanelixia albertana) was recorded as a confirmed identification. We also searched the
Consortium of North American Lichen Herbaria (https://lichenportal.org, CNALH 2020) for records, and
T. Spribille searched a draft modern catalog of lichens from the prairie provinces (Deneka et al. in prep.),
which includes a comprehensive summary of species cited in literature.
Data sources. III. Nature-reporting applications. – We searched two citizen science
applications (“apps”) for submitting photography-based observations of biodiversity, NatureLynx (2020)
and iNaturalist (2020), for lichen records within our geographic area. Data were downloaded directly from
iNaturalist using Edmonton as a geographic filter and a “fungi including lichens” taxon filter, providing
191 potential records. The geographic locations of observations uploaded to NatureLynx are masked for
privacy and protection of potentially sensitive data, so accurate location data was provided by the ABMI (J.
Bell, ABMI unpublished data, 2020). After filtering all provided records by location, 136 records from the
“bryophytes and lichens” category remained. All records were visually examined, records for bryophytes
and non-lichenized fungi were removed, and lichen photographic records were verified or reidentified
where possible, while also noting their substrate and location within Edmonton.
Identification. – We used Leica MZ6 or M60 stereo microscopes and Leica DM750 or Medilux
12 compound microscopes to examine anatomy under bright field or polarized light, noting the presence
(POL+) or absence (POL-) of birefringent crystals. Anatomical measurements are given as a range
(minimum–maximum) because of the small number of specimens available from Edmonton. Where
applicable we used chemical spot tests to support our determinations, including 10% potassium hydroxide
(K), sodium hypochlorite (C), Lugol’s iodine (I), paraphenylenediamine (PD, either dissolved in ethanol or
in the form of Steiner’s solution), ultraviolet light (either short wave [UV254] or long wave [UV365]), largely
following Brodo et al. (2001). Some collections were examined using thin-layer chromatography (TLC),
mostly following Orange et al. (2010). We used 10 × 20 cm glass plates with solvents A, B′, and C (Orange
et al. 2010), checked for fatty acids using water spray, and boiled the acetone-specimen mixtures three
times in a water bath prior to spotting (as per I. Brodo, pers. comm.). For photography, we used one of the
following: Leica MC170 HD camera (stereoscope), Leica ICC50 HD (compound microscope), Dinolite
Edge Digital Microscope AM4515Z7T, or a microscope-mounted Canon D6. We color-corrected and
created composite images from images taken at multiple focal depths to increase resolution in Photoshop
(Adobe 2008-2021). Sections of specimens were mounted in water unless otherwise indicated.
Molecular methods. – To verify or aid in the identification of a subset of collections, the internal
transcribed spacer (ITS ribosomal DNA; internal transcribed spacer regions 1 and 2 as well as the
embedded 5.8S region of the ribosomal rDNA and adjacent sections of the large and small ribosomal
subunits, LSU and SSU) was Sanger sequenced by T. Spribille’s lab at the U of A. ITS is the single most
sequenced locus in fungi and widely used as a barcode (Hoffman & Lendemer 2018, Schoch et al. 2012,).
DNA was extracted using the Qiagen DNeasy Plant Mini Kit following the manufacturer’s instructions, or,
in the case of sparse material, the QIAmp DNA Investigator Kit. PCR was performed using ITS1-F (Gardes
& Bruns 1993) and ITS4 primers (White et al. 1990), and the KAPA 3G Plant PCR Kit (KAPA
Biosystems). The PCR cycle used was: pre-denaturation for 5 min at 95°C, 35 cycles of amplification, each
cycle 30 sec at 95°C, 30 sec at 57°C, and 30 sec at 72°C. After the 35 cycles, extension occurred over 7
min of 72°C and then samples were stored at 4°C. PCR products were visualized on agarose gel after
electrophoresis and sent for sequencing if a product was seen. Samples with multiple bands were not sent
for sequencing due to poor chance of a clear sequence. Prior to sequencing, samples were purified using
standard ExoSap protocol. PCR products were sequenced by Psomagen, Inc., USA, and forward sequences
were visually examined for errors or ambiguities prior to screening.
Phylogenetic analyses. – We screened sequences with BLAST searches against the NCBI
nucleotide database to identify sequences that may represent non-target organisms (NCBI Resource
Coordinators 2018). The sequences generated for this study were complemented with sequences from
GenBank representing additional species and specimens, as well as a small number of sequences from the
senior author. For queried sequences of species adequately represented in GenBank, we report similarity
40
metrics with accessioned sequences in the annotated species list. Further analyses including de novo tree
construction were conducted for Candelariaceae, Endocarpon, Lecidella, Flavopunctelia, Lecanora
dispersa group, Peltigera, Physconia, and Punctelia, as BLAST results were insufficient.
For genera requiring phylogenetic analyses, the following steps were common across analyses;
specifics for each phylogeny are provided below. Sequences for each analysis were aligned with our query
sequence(s) using MAFFT via a web platform (MAFFT ver. 7.49, Katoh et al. 2002, Katoh & Standley
2013, Katoh et al. 2019) or in MegAlign Pro v. 17 (DNASTAR 2021), and visually inspected in BioEdit
7.7.1 (Hall 1999). We used ITSx 1.1 (Bengtsson-Palme et al. 2013) to split sequences into ITS, small
subunit, and large subunit files to aid in sequence vetting and where appropriate create partitions for
nucleotide substitution model fitting. We visually examined final alignments in BioEdit and trimmed all
sites from the alignment present in ≤10% of sequences. Alignments were screened using GUIDANCE2 for
ambiguous sites, and analyses were completed with and without ambiguous regions and the resultant trees
visually compared. Original fasta files and final alignments are deposited in Dryad (DOI
https://doi.org/10.5061/dryad.sqv9s4n6d). Sequence voucher data are provided in Supplementary Appendix
2, also available in the Dryad deposit. We generated maximum likelihood phylogenetic trees in W-IQ-
TREE 1.6.12 (Nguyen et al. 2015, Trifinopoulos et al. 2016) via http://iqtree.cibiv.univie.ac, specifying
partitions (partition model: Chernomor et al. 2016), linked branch lengths, automatic model selection
(ModelFinder: Kalyaanamoorthy et al. 2017), and free rate heterogeneity. Branch support was analyzed by
1,000 ultrafast bootstraps (UFBoot: Hoang et al. 2018) as well as SH-aLRT single branch tests with 1,000
replicates. Trees were visualized and organized in Dendroscope 3.7.6 (Huson & Scornavacca 2012) and/or
MegAlign Pro, and exported to Microsoft Office Professional Plus Powerpoint 2016 for editing.
The Candelariaceae phylogeny was generated de novo with seven new sequences from the senior
author, GenBank sequences with high BLAST similarity to our new sequences, and sequences from
Westberg et al. (2011), Liu & Hur (2018), and Liu et al. (2019). Additional sequences for Candelaria were
added from GenBank to increase taxon sampling in that clade. The Endocarpon phylogeny was generated
de novo using two new sequences from the senior author, GenBank sequences with high BLAST similarity
to our new sequences, and sequences from Zhang et al. (2017). The Flavopunctelia phylogeny was
constructed using all accessioned sequences of Flavopunctelia in GenBank, sequences from this study, and
GenBank sequences with high BLAST similarity to our new sequences, regardless of determination. The
Lecanora dispersa group phylogeny was created by adding new sequences from this study, their top-
scoring megablast GenBank sequences, and the ITS of the type of L. lendemeri E. Tripp & C.A. Morse
(Tripp et al. 2019) to the multiple sequence alignment from Śliwa et al. 2012 (Treebase study #12681,
using ‘mafft—add’ (https://mafft.cbrc.jp/alignment/server/add.html, Katoh & Frith 2012). Similarly, the
Physconia phylogeny was compiled using the 60 sequences from Esslinger et al. (2017, deposited in Dryad
as https://doi.org/10.5061/dryad.bh7mc), and additional sequences from the senior author, GenBank, and
this study. Finally, we aligned five new Punctelia sequences to the ITS portions of the concatenated
alignment of Alors et al. (2016), and the ITS alignment of Lendemer & Hodkinson (2010) using ‘mafft—
add’. ITS was concatenated with the other loci in Mesquite, and the new multiple sequence alignments
were reanalyzed with partitions.
For Peltigera sequences, we also used NCBI BLAST with megablast to check the percent of our
sequence that was identical to sequences published by F. Lutzoni and J. Miadlikowska Peltigera projects,
which we mapped to currently undescribed molecular species delimited by Pardo-De la Hoz et al. (2018)
and Magain et al. (2018). For Peltigera section Peltigera we also checked for the presence of species-
specific hypervariable region sequences described in Magain et al. (2018).
A sterile crust that could not be assigned to genus on morphology or chemistry alone (Haughland
2020-28) was analyzed using the workflow in Hodkinson & Lendemer (2012). We first queried the
sequence in NCBI BLAST with megablast. Based on the combination of the best BLAST hits, as well as
possible species matches from the literature based on TLC results (Lendemer 2010, 2013; Malíček et al.
2017), a multi-genus dataset was created to show the placement of the sequence within the potential genera
Buellia, Lecanora, Lecidella and Lepraria. Based on those results, we used the most recent Lecidella
phylogeny from Zhao et al. (2015; TreeBase Study ID #17997), downloaded the seven loci dataset, added
selected ITS sequences from GenBank and this study to the existing alignment (mafft--add, Katoh & Frith
2012), manually edited and trimmed the new alignment, and generated a new phylogenetic tree using
methods described above, with partitions for each locus. For the multi-genus tree, we post-hoc graphically
simplified and collapsed clades within Lecidella to focus on the broader, genus-level placement of our
sequence.
41
Nomenclature and new records. – Nomenclature and taxonomic authorities largely follow
Esslinger (2019). Exceptions are outlined at the beginning of each morphogroup section in the annotated
species list below, and for species for which we have adopted relatively recent taxonomic changes or
conversely retained older taxonomy we provide recent synonyms (e.g., names in Brodo et al. 2001). To
assess whether species were newly reported from Edmonton and Alberta, we searched salient literature, a
draft modern catalog of lichens for the prairie provinces (Deneka et al. in prep.), as well as the databases
previously mentioned (see section Data sources. II. Historical and herbarium records), using both currently
accepted nomenclature and synonyms.
Preliminary assessment of species’ utility as citizen science indicators. – In addition to being
responsive to the gradients of interest (air quality and climate), we decided a citizen science biomonitor
should be widely distributed across Edmonton, be easily perceptible and identifiable, and show high survey
repeatability. Here we address the latter; sensitivity to gradients will be addressed in a future paper.
Distribution was assessed using our composite dataset. We summarized species by the number of records
as well as their presence in each of three major habitat types: tablelands (largely represented by boulevard
trees), parklands (forested parks and natural areas within the tablelands), and river valley and ravine forests.
We used presence and abundance of records in nature-reporting apps as a metric of perceptibility to the
public. We compared the original, user-submitted identifications to our determinations to assess the ability
of the public to accurately identify lichens to genus or species. We assessed survey repeatability using
Pearson correlations and t-tests for significance between a species score on original surveys and audits at
50 sites for the 19 macrolichen species and 2 crustose genera included in the systematic surveys. We set a
cut-off of a statistically significant R≥0.5 to consider a surveyed species repeatable. Analyses were run in
Microsoft Office Professional Plus Excel 2016. Species maps were created using looping scripts in ESRI
ArcGIS Pro 9.2.0, based on a map created in ESRI ArcMap 10.7.1.
RESULTS
Phylogenetic analyses. – Our proximate goal was to confirm the taxonomic placement of a subset
of Edmonton collections. Of 44 collections for which DNA was extracted, we obtained sequences for 35
representing 33 putative species. Six collections were not a priori identified with confidence; sequencing
allowed us to at least tentatively identify four of those species. Of the remaining 27 species, molecular data
supported all of our phenotype identifications to genus, and 59% (16) of our identifications to species. Of
the remaining 41% (11), we could not confidently resolve the species-level taxonomy of 30% (8) due to
insufficient molecular data (lack of reference sequences, low BLAST values, and/or unresolved
polyphyletic species in phylogenetic analyses). We concluded our species-level identifications were
incorrect for 3 species (11%), and molecular data allowed us to correct those. In total we generated nine
phylogenies based on ITS, and these analyses provide some insight into the broader taxonomy of these
taxa, explored by taxon below.
Candelariaceae. – The first phylogeny for Candelariaceae and the nrITS sequences generated for
that study (Westberg et al. 2007) form the core of all subsequent analyses (e.g., Liu & Hur 2018,
Kondratyuk 2020, this study, Fig. 2). After alignment, trimming, and removal of ambiguous regions
(guidance score of <0.93), our dataset was composed of 86 sequences and 490 sites, of which 185 were
parsimony-informative, 64 were singletons, and 241 were constant. The best fit models by partition were
SSU: TN+F+G4, ITS1: GTR+F+G4, 5.8S: K2P, ITS2: TPM2+F+G4, and LSU: JC. The basal branches
remain poorly resolved, and some species as currently understood are poorly discriminated by ITS (e.g.,
Candelariella vitellina (Hoffm.) Müll. Arg). Given the need for additional loci and intra-specific
representation, here we do not propose taxonomic changes based on our analyses, instead highlighting
areas where revisions may be required in the future. One branch of the tree that has benefited from
additional data is Candelaria Clade III (following Westberg et al. 2007), including Candelaria concolor
(Dicks.) Arnold and the recently described Candelaria asiatica D. Liu & J.S. Hur (Liu & Hur 2018). Our
Alberta sequence clusters with a highly supported clade of C. concolor from eastern North America, sister
to a Candelaria asiatica clade (Fig 2). In contrast, C. concolor sequences from Europe form a highly
supported clade distinct from other regions (Fig. 2). Recent sequences of C. asiatica from China
(Kondratyuk et al. 2020) have not been deposited to a public repository, so geographic representation of
42
Figure 2. The maximum likelihood tree of Candelariaceae species based on nrITS. The tree was unrooted
for analyses, and rooted for visualization on Pycnora xanthococca. The numbers above each branch
represent the single branch support (%)/ultrafast bootstrap support (%); branches where both values ≥70%
are drawn with thicker lines. GenBank sequences are labelled by their accession number and collection
location. Newly-generated sequences are in bold. Scale = nucleotide substitutions per site.
43
Figure 3. The maximum likelihood tree of Endocarpon based on nrITS. The tree was unrooted for
analyses, and rooted for visualization on Verrucaria macrostoma. The numbers above each branch
represent the single branch support (%)/ultrafast bootstrap support (%); branches where both values exceed
70% are drawn with thicker lines. GenBank sequences are labelled by their accession number and name,
and monophyletic species are highlighted with grey polygons. Newly-generated sequences are in bold.
Scale = nucleotide substitutions per site.
44
Figure 4. The maximum likelihood tree of Flavopunctelia species based on nrITS. The tree was unrooted
for analyses, and rooted for visualization on Punctelia caseana. The numbers above each branch represent
the single branch support (%)/ultrafast bootstrap support (%); branches where both values exceed 70% are
drawn with thicker lines. GenBank sequences are labelled by their accession number and name. Newly-
generated sequences are in bold. Names in quotes are putative misidentifications in GenBank. Scale =
nucleotide substitutions per site.
that species within our analysis is limited. Kondratyuk et al. (2020) interpreted a single specimen from
Canada based on 28S nrLSU as C. asiatica; this sequence came from the same specimen as KT695365
(ITS used herein), but it appears that for unexplained reasons those authors did not use the corresponding
ITS sequence in their phylogeny.
Our sequences of what we morphologically classified as epiphytic Candelariella vitellina are
identical to the single available sequence of Candelariella efflorescens R.C. Harris & W.R. Buck, an
epiphytic, sorediate, rarely fertile species that occurs predominantly on broadleaved trees in eastern North
America (Harris & Buck 1978). Only additional data will differentiate between alternate interpretations of
this; either ITS is insufficient to discriminate species within this clade, or C. efflorescens is an epiphytic
species that varies phenotypically across its range. Harris and Buck (1978) noted in their description of C.
efflorescens that “given an isolated apothecium, we would not be able to distinguish which of the three
species [efflorescens, vitellina and xanthostigma (Ach.) Lettau] it came from.” Because our specimens were
never found to be sorediate, here we treat them as C. cf. vitellina pending further phylogenetic work.
Similarly, since our single C. lutella sequence did not cluster with existing sequences of C. lutella, and
instead was basal to the C. efflorescens clade, we treat those collections as C. cf. lutella.
Endocarpon. – After alignment and trimming, the final dataset consisted of 48 sequences and 848
sites, of which 263 were parsimony-informative and 71 were unique. The best fit models by partition were
SSU: K2P+G4, ITS1: TNe+G4, 5.8S: K2P+I, ITS2: TIMe+R2, and LSU: K2P+I. These analyses should be
interpreted cautiously as only 14 of the circa 75 currently accepted Endocarpon species are represented in
GenBank. Our single ITS sequence is basal to a highly supported clade of E. unifoliatum T. Zhang, X. L.
45
Wei & J. C. Wei recently described from China (Zhang et al. 2017), and here reported new to North
America (Fig. 3). In acknowledgement of the lack of representation of described species, as well as the
relatively large number of differences in our sequence versus the five sequences from China, we report our
determination as E. aff. unifoliatum.
Flavopunctelia. – Six Flavopunctelia species are accepted at present (Index Fungorum,
www.indexfungorum.org), and sequences exist for the three species reported to be sorediate: F. flaventior
(Stirton) Hale, F. soredica (Nyl.) Hale, and F. borrerioides Kurok. After alignment and trimming, the final
dataset was composed of 33 sequences and 511 sites, of which 40 were parsimony-informative and 123
were unique. No ambiguous regions were detected in guidance (alignment score=1.0), and the best fit
models by partition were ITS1: TNe, 5.8S: K2P, and ITS2: TNe+I by BIC (Kalyaanamoorthy et al. 2017).
This tree is the most comprehensive Flavopunctelia tree published to date, and overall supports the
separation of F. flaventior and F. soredica with high support (Fig. 4). These species often intergrade
phenotypically in Alberta. A single sequence of F. borrerioides groups in a highly supported clade with
two F. flaventior from Spain. Future work should re-examine these collections to determine if this clade
represents F. borrerioides s.str. Given we found no records of F. borrerioides in the CNALH, Consorcio de
Herbarios de Líquenes en América Latina (https://lichenportal.org/chlal/), or GBIF (https://www.gbif.org/),
additional work is needed to determine if this species is overlooked or whether it should be reduced to
synonymy with F. flaventior. Additional structure within F. flaventior is addressed in a submitted
publication (K.C, Rajeshkumar, B.O. Sharma and S. Fatima, pers. comm.). Our single ITS sequence of F.
soredica is basal to a well-supported clade of F. soredica sequences. Three sequences from other genera
with high BLAST similarity to our sequence may be misidentifications (denoted by quotes within the tree).
Lecanora dispersa group – Given the recent flux in generic designations within Lecanora (Zhao et
al. 2016, Kondratyuk et al. 2019), here we retain Lecanora in the broad sense. After alignment and
trimming, the final dataset was composed of 66 sequences and 511 sites, of which 180 were parsimony-
informative and 56 were unique. The best fit models by partition were ITS1: TIM2+F+G, 5.8S: K2P, and
ITS2: TNe+G4 according to BIC (Kalyaanamoorthy et al. 2017). The final maximum likelihood tree
suggests many of the same relationships as in Śliwa et al. (2012) and Tripp et al. (2019), and also presents
the same unresolved problems (Fig. 5). Our principal concern was the placement of L. hagenii (Ach.) Ach.
and L. cf. persimilis (Th. Fr.) Arnold sequences from Edmonton. It is clear L. hagenii as currently
understood is polyphyletic. It is not clear whether any of the four clades containing L. hagenii sequences
correspond to that species in the strict sense. Śliwa et al. (2012) hypothesized that the clade marked
“hagenii II” on our tree may correspond to hagenii s.str., due to its close relationship to L. crenulata
Hooker (Fig. 5). This clade is widely distributed and contains both pruinose and epruinose collections,
including our L. cf. persimilis. Conversely, our single Edmonton sequence of L. hagenii clusters in a new,
highly supported clade of North American pruinose specimens (“hagenii III”; Fig. 5), outside of all prior
hagenii clades (numbered I, II, and IV). In summary, the analyses confirmed only that our sequences
cluster within the highly supported L. dispersa group, and that the sequences represent distinct taxonomic
entities. Without further resolution, we acknowledge the uncertainty by modifying both epithets with “cf.”
Lecidella. – A specimen that could not be identified without molecular data (isolate DLH1 from
Haughland 2020-28) was first analyzed to determine its generic affinities using sequences of
phenotypically similar species from different genera (Fig. 6B). Species we considered in addition to
matches generated in BLAST include Buellia arborea Coppins & Tønsberg, Lecanora alboflavida Taylor,
L. allophona (Ach.) Nyl., L. impudens Degel., and Lepraria rigidula (B. de Lesd.) Tønsberg. Other
potential sterile sorediate/granular species that we did not find ITS data for (Cliostomum, Lecanora,
Lepraria, and Rinodina) could be excluded by TLC results (our specimen contained atranorin and two
possible xanthones that fluoresce orange under UV365 light). After BLAST, sequence selection, alignment,
and removal of ambiguous sites (guidance score <0.93), our tree-building alignment was composed of 48
sequences and 421 sites, of which 105 were parsimony-informative, 110 were singletons, and 206 were
constant. The best fit model was TNe+R2 using BIC. Analyses suggested our sequence fell within
Lecidella, close to a single GenBank sequence of Lecidella albida Hafellner (Fig. 6B). The second analysis
focused on placement within Lecidella, contained 78 sequences and 4084 sites, of which 1460 were
parsimony informative, 1828 were singletons, and 796 were constant. The best fit models by BIC for each
locus were: TIM2+F+G4 for ITS, TNe+R4 for nuclear ribosomal large subunit (LSU), TN+F+R2 for
46
Figure 5. The maximum likelihood tree of the Lecanora dispersa group based on nrITS. The tree was
unrooted for analyses, and rooted for visualization on Lecanora allophana. The numbers above each
branch represent the single branch support (%)/ultrafast bootstrap support (%); branches with both values
≥70% are drawn with thicker lines. GenBank sequences are labelled by their accession number and specific
epithet. Newly-segregated genera previously within Lecanora are labelled as Lecanora regardless of
GenBank genus for ease of comparison. Monophyletic species are highlighted with grey polygons, and
polyphyletic clades of L. “hagenii” are outlined in empty polygons. Newly-generated sequences are in
bold, and sequences new to the Śliwa et al. 2012 dataset are prefaced with ‘’. Scale = nucleotide
substitutions per site.
47
Figure 6. The maximum likelihood trees showing (A) placement of Edmonton sequences within Lecidella based on
Zhao et al. (2015) and (B) placement of Edmonton sequences within a range of phenotypically-similar yet
phylogenetically-distant species, based on nrITS. The trees were unrooted for analyses, and rooted for visualization
with (A) Rhizoplaca porteri and (B) Lepraria membranacea. Values above each branch represent the single branch
support (%)/ultrafast bootstrap support (%); branches where both values exceed 70% are drawn with thicker lines.
GenBank sequences are labelled by their ITS accession number and name. Newly-generated sequences are in bold, and
sequences new to the Zhao et al. 2015 dataset are prefaced with ‘’. In tree B, the species within the Lecidella
elaeochroma and L. stigmatea clade are collapsed to emphasize the deeper phylogenetic structure. Scale = substitutions
per site.
48
Figure 7. The maximum likelihood tree of Peltigera species based on the nrITS. The tree was unrooted for
analyses, and rooted for visualization on Peltigera rufescens. The numbers above each branch represent the
single branch support (%)/ultrafast bootstrap support (%); branches where both values exceed 70% are
drawn with thicker lines. GenBank sequences are labelled by their ITS accession number and name.
Newly-generated sequences are in bold. Scale = nucleotide substitutions per site.
mitochondrial small subunit (mtSSU), TNe+G4 for minichromosome maintenance complex component 7
(MCM7), TN+F+R3 for the largest subunit of the RNA polymerase II gene (RPB1), K2P+G4 for the
second largest subunit of RNA polymerase II gene (RPB2), and K2P for ribosome biogenesis gene (TSR1).
Similarly, our sequence formed a highly supported clade with a single sequence of Lecidella albida, distant
from other sequenced Lecidella species (Fig. 6A). Lecidella albida is apparently new to North America.
Future analyses should consider whether L. albida belongs outside of Lecidella s.str. and should be treated
as distinct.
The Lecidella clades from Zhao et al. (2015) were also recovered in our analyses, with our
additional sequences largely forming their own branches within those clades. Lecidella elaeochroma (Ach.)
M. Choisy was recovered as polyphyletic, and our second Lecidella sequence, originally determined as L.
euphorea (Flörke) Kremp. (isolate DLH11 from Haughland 2020-43), is positioned on a well-supported
branch separate from but close to a clade with L. “elaeochroma 5” from Europe (Zhao et al. 2015). Because
of this relationship, we amended the identification of our L. euphorea to L. elaeochroma. We anticipate our
taxonomy will require further revision, particularly given Lendemer et al. (2019) concluded that at least
some of the type material of L. elaeochroma does not match the material to which the name is currently
being applied. Much more work is required to untangle Lecidella.
Peltigera. – Molecular data confirmed the identity of eight of the 11 Peltigera species found in
Edmonton (Fig. 7). It also reinforced the challenge of accurately identifying members of Peltigera section
Peltigera. For example, a specimen originally identified as P. membranacea (Ach.) Nyl. was recovered
within the P. praetextata clade; the latter species shows considerable phenotypic plasticity, even within
localities in close proximity, growing in common urban conditions. We formally report P. islandica
Goward & S.S. Manoharan-Basil as new to Alberta, previously documented within Alberta by the ABMI
(unpublished, in collaboration with C. Pardo-De la Hoz, F. Lutzoni, J. Miadlikowska, T. Goward, and I.
Medeiros).
49
Figure 8. The maximum likelihood tree of Physconia species based on the nrITS, and the phylogeny in Esslinger et al.
(2017). The tree was unrooted for analyses, and rooted for visualization on Anaptychia elbursiana. The numbers above
each branch represent the single branch support (%)/ultrafast bootstrap support (%); branches where both values exceed
70% are drawn with thicker lines. GenBank sequences are labelled by their ITS accession number and name. Newly-
generated sequences from Edmonton are in bold, and sequences new to the Esslinger et al. dataset are prefaced with
‘’. Scale = nucleotide substitutions per site.
50
Figure 9. The maximum likelihood tree of clade C from the addition and reanalysis of Alors et al. (2016),
based on the nrITS. The numbers above each branch represent the single branch support (%)/ultrafast
bootstrap support (%); branches where both values exceed 70% are drawn with thicker lines. Sequences are
labelled following the TreeBase supplementary data of Alors et al., sequences new to the Alors et al.
dataset are prefaced with ‘’and the newly-generated sequence from Edmonton is in bold. Scale =
nucleotide substitutions per site.
Physconia. – After alignment and trimming, the final dataset consisted of 86 sequences and 486
sites, of which 111 were parsimony-informative and 43 were unique. Sequences included 60 from Esslinger
et al. (2017), two from Edmonton, 11 from northern Canada from the senior author, and 13 additional
GenBank sequences chosen for their high similarity to new sequences or to represent species missing in the
original tree. Of the 14 species known from North America (Esslinger 2019), ten were well-represented in
Esslinger et al. (2017), and we added sequences of P. isidiomuscigena Essl. to represent 11 North
American species in total. The three species missing sequence data are the esorediate P. californica Essl.,
the fertile and lobulate P. subpallida Essl., and P. fallax Essl., which forms “nest-like” soralia; none of
these species are likely to be confused with our sorediate collections. Globally, approximately 31
Physconia species are recognized (depending on which synonymies are accepted), 19 of which are
represented in our analyses (Fig. 8). The best fit models by partition were ITS1: TIM2e+G4, 5.8S:
K2P+I+G4, ITS2: TNe+G4. The analyses support our identification of P. detersa and P. enteroxantha, with
the latter nesting within a well-supported monophyletic clade (Fig. 8). A single, newly added specimen of
P. jacutica Urbanav., Ahti & Loht. from GenBank nests within an otherwise monophyletic clade of P.
detersa. Most Physconia species form well-supported monophyletic clades. A noteable exception is P.
muscigena (Ach.) Poelt. In addition, some sequences of specimens originally identified as P. perisidiosa
(Erichsen) Moberg from northern Canada cluster with P. rossica Urbanav., known only from Russia and
China prior to these analyses. Re-examination of these specimens suggest that they fit within the
morphological circumscription of P. rossica. Additional work is ongoing with T. Esslinger and S. Leavitt
on the Physciaceae.
51
Data source
# collections
examined
# species
confirmed
# unique
records
# unique first
AB records
Opportunistic surveys
(2013–present)
180
89
26
3 + 2 putative
undescribed
species
Herbarium historical records
(1974–1988)
129
47
3
0
U of A deciduous epiphytes
grid-based surveys
>5000 observations,
191 mixed collections
43
19
3
U of A rare tree surveys
190 observations, 44
mixed collections
23
1
0
Nature-reporting app submissions
195
47
5
1
NatureLynx
(128)
(40)
(3)
(1)
iNaturalist
(67)
(20)
(0)
(0)
Literature reports
N/A
19
0
N/A
Table 1. Species detected by data source and their relative contribution to unique (not found in any other
source) or new species records.
Punctelia. – Our sequences grouped with Punctelia caseana in both our re-analysis of Lendemer
& Hodkinson (2010, isolate DLH10, results not shown), and Alors et al. (2016). Here we show just the
branch of the phylogeny with P. caseana and P. jeckeri (Clade C in Alors et al. 2016) to support our
identification of P. caseana (Fig. 9).
Additional taxa with molecular data. – We provide sequence comparative data within the
annotated species list for the following species: Bacidia circumspecta (Nyl. ex Vain.) Malme,
Blennothallia crispa (Hudson) Otálora, P. M. Jørg. & Wedin, Caloplaca feracissima H. Magn., C. tominii
(Savicz) Ahlner, Lecania naegelii (Hepp) Diederich & van den Boom, and Lepraria finkii (B. de Lesd.) R.
C. Harris. Analyses for these sequences were restricted to BLAST comparisons due to their high percent
identity, BLAST scores and query coverage with multiple accessioned sequences that matched our original,
phenotypic determination. The Edmonton sequence of Ramalina pollinaria (Westr.) Ach. is analyzed in
Haugland et al. (in prep.), and it forms a monophyletic clade with previously published sequences of R.
pollinaria s.str.
Diversity. – We found herbarium or literature records for 53 lichens within Edmonton. The
species recorded in nature-reporting apps, largely generated over the last five years, contributed
observations of 17 additional species, for a total species richness of 70. With field work, we were able to
verify and find extant populations for all but five of those 70 species, and we generated records for an
additional 44 species (largely crustose, calicioid or allied fungi). In total, we documented 133 species from
across all data sources, including >620 collections and >5,000 field observations (Table 1). Below, we
present an annotated list for 114 species (see also Supplementary Appendix 1). Of the remaining 18
species, seven represented misidentifications (Bryoria fremontii (Tuck.) Brodo & D. Hawksw., B. glabra
(Motyka) Brodo & D. Hawksw., Cladonia ecmocyna Leighton, Peltigera aphthosa (L.) Willd.,
Phaeophyscia hispidula (Ach.) Essl., Physcia millegrana Degel., and Usnea dasopoga (Ach.) Nyl.). Data
from the ABMI’s systematic, province-wide surveys suggests these species are not likely found in
Edmonton because a) their distribution is largely limited to the Foothills and/or Montane Natural Regions
of Alberta (B. fremontii, B. glabra, C. ecmocyna) or to the cooler Boreal region (Peltigera aphthosa), or b)
they are very rare, restricted in their distribution, or even absent from Alberta (Phaeophyscia hispidula,
52
Figure 10. Proportion of lichens by a) growth forms and b) photobiont classifications detected in the city of
Edmonton in comparison to those known for the province of Alberta (980 species, Government of Alberta
2017).
Physcia millegrana). Another three species records were based on outdated taxonomy (Caloplaca
holocarpa (Hoffm. ex Ach.) A.E. Wade, Physconia grisea (Lam.) Zahlbr., and Ramalina fastigiata (Pers.)
Ach.) and were revised.
We had information supporting the presence of an additional six species but were not able to
confirm those records; we consider these open investigations and will continue to seek specimens to
corroborate their presence. They include Pseudevernia consocians (Vain.) Hale & W. L. Culb.,
Melanohalea subolivacea (Nyl.) O. Blanco et al., and Micarea melaena (Nyl.) Hedl., which were based on
herbarium collections catalogued in BIOTICS (Government of Alberta 2020) or PMAE. Unfortunately,
these specimens are missing, even after herbaria searches by us or colleagues at ALTA (T. Spribille and C.
La Farge-England, pers. comm.) under similar species and synonyms. In addition, Ochrolechia arborea
(Kreyer) Almb. was reported by Elsinger et al. (2007), and Brodo (1991) cited a collection of this species
from a protected area west of Edmonton, but we could not verify its presence within Edmonton. Two
species we could not confirm fit multiple categories. Collections of putative Cladonia rei Schaerer and
Usnea glabrata (Ach.) Vain. were redetermined to other species, but geographically and ecologically it is
possible that they will yet be found within Edmonton (ABMI 2020, Haughland et al. 2018). Finally, we
have specimens representing four species (Caloplaca sp., Candelariella xanthostigma, cf. Lepra sp.,
Rinodina cf. albertana Sheard) that are too sparse or poorly developed to present with any confidence at
this time.
Species richness was highest in foliose lichens (48 species), followed by crustose, calicioid and
allied fungi (41 species), and lowest in fruticose lichens (25). Edmonton’s lichen flora is largely
representative of that of Alberta with some notable divergences. Crustose lichens, fruticose lichens,
cephalodiate lichens, and epiphytic cyanolichens were under-represented in Edmonton’s flora (Figs. 10 and
11), while we documented a surprising number of terricolous cyanolichens, almost exclusively Peltigera.
No epiphytic Leptogium, Lobaria, or Nephroma species were detected, despite searches of large-diameter
trees in mixedwood and deciduous riparian forests that commonly house these genera in the Boreal Natural
Region surrounding Edmonton. Finally, we found only three Collemataceae species in Edmonton, and
Cladoniaceae are under-represented relative to their high diversity in the province (Fig. 11).
Opportunistic surveys resulted in the greatest number of species records with the smallest overall
effort, but the U of A systematic and rare tree surveys provided a similar number of unique records, largely
of crustose species (Table 1). PMAE specimens were representative of the macrolichen species from
Edmonton, but few crustose lichens were represented.
New records. – In addition to the two putative new records for North America (Endocarpon aff.
unifoliatum and Lecidella albida), we report an additional twelve species new to Alberta: Amandinea
dakotensis (H. Magn.) P. May & Sheard, Bacidia circumspecta, Candelaria pacifica, Candelariella
53
Figure 11. Comparison of taxonomic families represented in Edmonton’s flora (23 different families) in
comparison to Alberta (58 families: Government of Alberta 2017; Lücking et al. 2017a, b). Only the top ten
families are listed, the remainder contribute <2% of the species each. Families are organized by highest to
lowest percentage and the colors of the top 10 families are consistent across charts.
antennaria Räsänen, Heterodermia japonica (M. Satô) Swinscow & Krog., Lecania naegelii, Lecanora
sambuci (Pers.) Nyl., L. stanislai Guzow-Krzemińska, Łubek, Malíček & Kukwa, and Lecidea
erythrophaea Flörke ex Sommerf., Peltigera islandica, Phaeocalicium aff. tremulicola (Norrlin ex Nyl.)
Tibell, and Xanthoria parietina (L.) Th. Fr. (Table 1, Supplementary Appendix 1, annotated list below).
The ABMI has publicly accessible records of the macrolichens Candelaria pacifica, Heterodermia
japonica, and Peltigera islandica, dating back to 2003; however, ours are the first published reports of
these species for Alberta. Five of the crustose species new to Alberta are apparently rare and were collected
almost exclusively from riparian and ravine habitats. In comparison, L. sambuci and C. antennaria were
collected largely from tableland habitats, and the latter species is surprisingly common considering it has
evaded detection until now.
In addition, two putative new species to science are reported for the genus Phaeocalicium, and
collections that could not confidently be attributed to a known North American species are reported from
Caloplaca s.l. and Lichinaceae. Molecular work on the calicioid lichens and allied fungi is underway in
collaboration with S. Selva and T. McMullin. Here we present a preliminary description to alert others to
these potential new species so that additional records may be sought. We also report records of Physcia
collections traditionally placed in existing species, but which ongoing molecular analyses suggest should be
separated as new species to science: Physcia aff. dimidiata (Arnold) Nyl. and Physcia aff. stellaris (L.)
Nyl. We defer further discussion of those taxonomic novelties to a future publication with S. Leavitt and T.
Esslinger. We also provide modern day reports of Alyxoria varia (Pers.) Ertz & Tehler for Alberta.
Previous records (Raup 1928, 1930) are from the northern Canadian Shield Natural Region in the province.
There may be a historical collection of A. varia from Edmonton in PMAE (under Opegrapha varia),
however the specimen is currently missing.
Assessment of species utility as citizen science indicators. – Approximately 25% (27 spp.)
detected occur in two or more of the three major habitat types, and another 13% (14 spp.) were detected
exclusively in the dominant tablelands habitats. Of those 41 species, we estimate 29 were widely
distributed and detectable enough to include in a preliminary target lichen survey list (Supplementary
Appendix 1). In comparison, 62% (68 spp.) were detected solely in the highly restricted river valley and
ravine habitats. The nature-reporting apps suggest that the most perceptible genus to the public was
Peltigera, followed by Xanthomendoza and Cladonia (Table 2).
The accuracy of genus-level identification was high overall. However, accuracy was lower for
genera of species in Alberta that are morphologically similar to common genera from eastern North
America that often dominate records in the apps, but are rare or absent from Alberta (e.g., Flavopunctelia
was commonly misidentified as Flavoparmelia caperata (L.) Hale in Alberta). Species-level identification
54
Genus
User Identification
Correct
Total #
Sightings
Brown foliose (Melanelixia, Phaeophyscia, Physconia)
100%
6
Cladonia
100%
16
Flavopunctelia
47%
15
Parmelia
60%
15
Peltigera
98%
49
Physcia
85%
13
Ramalina
100%
6
Xanthoria s.l. (Rusavskia, Xanthomendoza)
75%
28
All other genera (Candelaria, Candelariella, Chaenotheca,
Punctelia, Hypogymnia, Lepraria, Vulpicida)
88%
8
Not verifiable
N/A
16
Total
78%
185
Species
User Identification
Correct
Total #
Sightings
Cladonia spp.
53%
15
Flavopunctelia flaventior
45%
11
Fruticose species (Bryoria fuscescens, Evernia mesomorpha,
Usnea spp.)
17%
6
Parmelia sulcata
47%
15
Peltigera canina group
47%
17
Peltigera elisabethae
44%
9
Other Peltigera (didactyla, leucophlebia, neckeri)
50%
4
Physcia adscendens, P. aipolia group, P. aff. dimidiata, P. aff.
stellaris
73%
11
Ramalina pollinaria
100%
5
Xanthomendoza fallax
50%
22
All other species
75%
16
Not identifiable
N/A
19
Total
47%
150
Table 2. Accuracy of user-submitted identifications at genus and species level, for taxa with ≥5 records
combined across NatureLynx and iNaturalist. For comparison, 58% (75 of 129) of PMAE specimens were
considered accurate at species-level using current taxonomy. Entries in bold indicate ≥80% accuracy.
accuracy was lower (47% for species vs. 78% for genera) but was similar to or exceeded PMAE overall
accuracy for Cladonia spp., Physcia spp., Ramalina pollinaria, and an assortment of species with few
reports (Table 2). Audits of U of A surveys found that seven of the 21 species were too rare to assess, seven
species met our cut-off of R≤0.5, and seven species had high inter-observer variability, and failed to meet
our cut-off (Table 3). While rare, the presumably sensitive fruticose lichens met that cut-off, both
individually or lumped together after the survey. Usnea specimens were often too small and poorly
developed to be identified to species. Of the foliose lichens, the three most common species showed high
repeatability (Xanthomendoza fallax, Phaeophyscia orbicularis, and Physcia adscendens; Table 3). Physcia
was otherwise a difficult genus for student surveyors to differentiate in the field: while almost all species
were under-detected by students, P. aff. dimidiata was over-detected, and P. aipolia group and P. aff.
stellaris were often confounded. For the latter two species, post-hoc lumping increased survey repeatability
to our cut-off (Table 3). Of the two crustose genera added to the target list, Caloplaca met our repeatability
55
Original species list
# Original/Audit
sites detected
R
T
P
*Caloplaca spp.
17/46
0.506
4.06
<0.001
Candelaria concolor/pacifica
5/13
0.371
2.77
0.008
Evernia mesomorpha
2/1
0.700
6.79
<0.001
*Candelariella spp.
3/47
0.174
1.23
0.226
Flavopunctelia flaventior
11/14
0.684
6.50
<0.001
Hypogymnia physodes
0/0
Too rare to assess
Melanelixia albertana
0/2
Too rare to assess
Melanelixia subaurifera
0/0
Too rare to assess
Melanohalea exasperatula
0/1
Too rare to assess
Parmelia sulcata
1/4
-0.042
0.29
0.771
Phaeophyscia orbicularis
50/50
0.757
8.03
<0.001
Physcia adscendens
44/48
0.555
4.63
<0.001
Physcia aipolia group
16/16
0.197
1.40
0.169
Physcia aff. dimidiata
17/6
0.171
1.20
0.236
Physcia aff. stellaris
29/41
0.409
3.10
0.003
Physconia spp.
0/1
Too rare to assess
Punctelia caseana
0/0
Too rare to assess
Usnea spp.
3/4
0.757
8.03
<0.001
Vulpicida pinastri
1/0
Too rare to assess
Xanthomendoza fallax
50/50
0.864
11.91
<0.001
Xanthomendoza hasseana
11/22
0.337
2.48
0.017
Post-hoc groups
# Original/Audit
sites detected
R
T
P
Physcia aff. stellaris/
P. aipolia group
39/42
0.624
5.53
<0.001
Usnea/Evernia
3/4
0.757
8.03
<0.001
Table 3. Repeatability of species assessments for lichens on the original survey list, as assessed with
Pearson Correlations between the original and audited values at 50 sites. An asterisk (*) marks crustose
genera added late to the survey list. Significance of the correlations was determined with a two-tailed T-test
(df=48, column T). Post-hoc groupings were calculated as the maximum value (from 1–4) observed of the
species in the group at each tree. Bolded entries indicate species with statistically significant R>0.5
(column R).
criterion. We summarize the Edmonton species that we feel meet enough criteria to form a preliminary
target list of potential lichen indicator species (Table 4).
DISCUSSION
Our primary goals were to address the deficit of lichen knowledge for Edmonton and to document
the species available for citizen science monitoring by publishing the first annotated lichen list for the city.
We doubled the known lichen richness, largely through systematic and opportunistic field work,
complemented by observations in nature-reporting apps. We also added 14 species to our provincial flora,
with the potential for additional species that are putative new species to science or currently evade
classification. This first annotated list presents data for 114 species within Edmonton, but we anticipate
many more species remain to be discovered. Saxicolous, conifer-dwelling, and crustose lichens require
additional survey effort. Leprarioid taxa require significantly more study in our region.
56
Species/Group
Growth
form
Photo-
biont
Veg.
Repro.
Habitats
Obs.
Chall-
enges
1
Caliciopsis calicioides
calicioid
n/a
none
P
1
det?
2
Phaeocalicium populneum
calicioid
n/a
none
R
P
7
det?
3
Caloplaca pyracea
crustose
chl
none
R
P
T
58
4
Caloplaca cerina
crustose
chl
none
R
P
T
5
det?
5
Candelariella antennaria
crustose
chl
none
T
6
6
Candelariella cf.
vitellina/lutella
crustose
chl
none
R
P
T
37
7
Lecanora impudens
crustose
chl
sor
R
P
8
8
Lecanora dispersa group
crustose
chl
none
R
P
T
14
det?
9
Rinodina spp.
crustose
chl
none
T
57
det?
10
Flavopunctelia spp.
foliose
chl
sor
R
(P)
(T)
>50
11
Melanelixia albertana
foliose
chl
sor
R
P
>10
12
Melanohalea exasperatula
foliose
chl
isi
R
(T)
5
13
Parmelia sulcata
foliose
chl
sor
R
P
(T)
15
14
Peltigera spp.
foliose
cya
varied
R
(T)
>28
15
Phaeophyscia kairamoi
foliose
chl
sor
R
P
3
16
Phaeophyscia nigricans
foliose
chl
sor
R
P
T
15
det
17
Phaeophyscia orbicularis
foliose
chl
sor
R
P
T
>191
18
Physcia adscendens
foliose
chl
sor
R
P
T
183
19
Physcia aipolia group
foliose
chl
none
R
P
T
61
id
20
Physcia dimidiata
foliose
chl
sor
R
P
(T)
19
id
21
Physcia aff. stellaris
foliose
chl
none
(R)
P
T
>157
id
22
Vulpicida pinastri
foliose
chl
sor
R
3
23
Xanthomendoza fallax
foliose
chl
sor
R
P
T
>191
24
Evernia mesomorpha
fruticose
chl
sor_isi
R
P
(T)
5
25
Ramalina spp.
fruticose
chl
most
sor
R
(T)
>7
26
Usnea spp.
fruticose
chl
sor_isi
R
(P)
(T)
>9
Table 4. Proposed preliminary epiphytic lichen list for citizen science biomonitoring in Edmonton based on
detectability, distribution across different habitats, and accuracy of identification. Photobiont:
chl=chlorolichen, cya=cyanolichen, n/a=non-lichenized calicioid fungus. Veg. Repro.=vegetative
reproduction; sor=sorediate, isi=isidiate. Habitats: R=river valley and ravines, P=parkland forests on the
flat tablelands, T=tableland habitats, largely boulevard trees and anthropogenic substrates. Parentheses
indicate the species/group is rare in that habitat. Obs.=number of observations across the various sources.
Challenges: det=low detectability, id=identification accuracy below desired cutoff, requires additional
training.
We have not focused on species currently considered provincially rare as the Alberta NatureServe
ranks require updating. However, we highlight a calicioid species found in Edmonton that our experience
suggests is truly rare and deserving of protection: Chaenotheca stemonea (Ach.) Müll. Arg. (ranked S1,
rare and tracked, Government of Alberta 2017) was a surprising find and is only the sixth collection known
to the senior author from Alberta. It was collected from a humid riparian zone with dense vegetation and
large diameter Picea (living and dead), where we also recorded two other calicioids and very high Peltigera
diversity. These habitats seem to be critical reservoirs of lichen diversity within the otherwise dry
tablelands upon which the majority of Edmonton lies. The importance of these riparian habitats is further
emphasized when we consider that while they occupy only 10% of the study area, 85% of the species
confirmed in this study were found at least once in a river valley or ravine habitat, and 62% were found
there exclusively.
57
Edmonton’s current lichen richness is comparable to estimates of urban lichen richness elsewhere,
including New York City, U.S.A (103 spp., Allen 2020), Geneva, Switzerland (127 spp., Habashi & Clerc
2013), and Grenoble, France (83 spp., Gombert et al. 2004). However, it pales in comparison to
municipalities in southern Ontario, one of Canada’s most densely populated and climatically mild regions.
While inclusive of relatively large, intact natural areas, a study of Toronto lists 180 species (McMullin et
al. 2019), and 543 species are now documented within a 50 km radius of Ottawa (Brodo 1988, Brodo et al.
2021b). Many studies that report lower species richness are restricted to epiphytes, making comparisons
between species lists difficult. However, Edmonton’s epiphytic calicioid lichen and allied fungi richness
appears to be high (nine species versus a mode of one in other studies; notable exceptions are four species
in Coxson et al. [2014] and 11 in McMullin et al. [2019]); this may be due to the interest of the authors in
calicioids or these urban areas could be genuine biological hotspots. Calicioid ecology and diversity are
poorly known in the continental interior of North America.
Studies from different cities in diverse jurisdictions including eastern Canada (Ontario, Nova
Scotia), Brazil, Spain, Sri Lanka, and the eastern United States continue to support the use of lichens as air
quality bioindicators (Koch et al. 2016, 2019; McCarthy et al. 2009; McMullin et al. 2017, 2019; Sergio et
al. 2016; Tulumello 2010; Will-Wolf et al. 2015; Yatawara & Dayanada 2019). Urban areas with poor air
quality were found to have low lichen species diversity (Coffey & Fahrig 2012, Koch et al. 2016, 2019,
McCarthy et al. 2009, Stringer & Stringer 1974, Yatawara & Dayanada 2019). Lichen recolonization
observed in conjunction with improvements in air quality in recent decades is further evidence of this
relationship (e.g., Allen 2020, Rose & Hawksworth 1981, Seaward & Letrouit-Galinou 1991). Urban areas
tend to be dominated by nitrophytic lichen species such as Candelaria concolor and Physcia spp. that are
less sensitive to industrialization (McCarthy et al. 2009, Sergio et al. 2016, Tulumello 2010).
Comparatively, these species are less abundant in areas further from urbanization with higher air quality,
where sensitive, fruticose species such as Usnea and Ramalina are present (McCarthy et al. 2009, Sergio et
al. 2016). Koch et al. (2019) found that cyanolichens, relatively loosely attached lichens, isidiate lichens,
and lichens partnered with Trentepholia indicated areas of low urbanization and low contaminant levels,
while chlorococcoid algae, narrow-lobed foliose species, sorediate species, and pruinose thalli were
indicators of medium-to-high urbanization and contaminant levels. The latter describes well the lichens that
dominate Edmonton’s tablelands.
However, researchers have long realized that patterns of urban lichen diversity are not shaped by
air quality alone (e.g., Brodo et al. 2021b, Golubkova & Malysheva 1978, Skye 1968). Urban areas tend to
be drier and warmer than rural areas, further lowering lichen diversity and abundance (McMullin et al.
2016, Yatawara & Dayanada 2019). The urban heat island was hypothesized to contribute to the lack of
lichens in downtown Winnipeg, Manitoba (termed a lichen desert by Stringer & Stringer 1974) given the
area’s low estimated sulphur dioxide. Even in the face of high sulphur levels, the tolerance of some species
to sulphur dioxide in dry, continental climates may be higher than in more oceanic climates (Hawksworth
et al. 1973). This suggests that complex interactions between climate, substrate, and pollutants should be
considered in future studies of urban lichen diversity. Future analyses of the data presented herein will
attempt to parse out the impacts of climate versus air quality in shaping species’ realized niche.
Survey limitations & directions for future research. – When interpreting these results, it is
important to recognize the limitations inherent in each data source. Survey effort is unequal across
substrates and habitats, and systematic surveys are restricted to deciduous trees. Rocks and similar
anthropogenic substrates such as buildings, sidewalks, walls, and cemeteries are particularly poorly
represented here. Downed wood and conifers also are under-surveyed, but to a lesser extent as they have
been better addressed through opportunistic surveys by the senior author and rare tree surveys in 2021.
Opportunistic surveys are biased towards naturalized areas and parks, as are nature application reports.
Absences cannot be inferred from locations with opportunistic, nature app, and herbarium records because
of unquantified survey effort. Crustose lichens were not collected at all systematic biomonitoring sites for
laboratory identification. Future gradient analyses will likely be restricted to a subset of sites where
samples were taken or for the city as a whole, and only to genus-level or morphological grouping. This
limitation affects our understanding of the distribution of those species, so maps presented herein
underestimate the niche and range of some species. Future surveys focused on under-sampled substrates
may remedy these issues and help form a more comprehensive view of diversity within the city.
With a minimum estimated error rate of 11%, our limited molecular analyses show the value of
barcode-level analyses in increasing the accuracy of determinations. It also suggests that despite our best
58
efforts, identification errors remain in the list presented here. Alberta in general would benefit from
inclusion in more taxonomic treatments. Lichen taxonomic treatments in North America often have either
an eastern or western focus, with relatively few collections from the continental interior where the two
floras intersect (e.g., Halonen et al. 1998, Lendemer 2009). In Alberta, we have documented species
previously thought to be restricted to eastern North America (e.g., Cladonia robbinsii A. Evans,
Pseudevernia consocians [ABMI 2020]), as well as species otherwise largely restricted to west of the
Rocky Mountains (e.g., Seirophora contortuplicata, (Ach.) Frödén, Hypogymnia imshaugii Krog,
Xanthomendoza montana (L. Lindblom) Søchting, Kärnefelt & S.Y. Kondr. [ABMI 2020, Haughland,
unpub.]). The paucity of molecular work in the interior and the intersection of eastern and western species
means that we are challenged to discriminate morphologically similar species that elsewhere can be
discriminated based on geographic range. Our expectation is that these species should be identified
accurately to properly understand their status, niche, and sensitivity. To do so will require more molecular
and morphological study in the quest of high-fidelity diagnostic traits.
With these limitations in mind, we make suggestions regarding which taxa should be investigated
further as urban biomonitors.
Suggestions for future biomonitoring. – Across macrolichen growth forms, genera were more
accurately identified than species in nature apps (Table 2) and by trained novices (Table 3). Genera that
were recognized, identified accurately, and met our repeatability cut-off include Flavopunctelia (if we a
priori exclude Flavoparmelia caperata), fruticose genera (Evernia, Usnea), and Physcia. Genera that are
relatively abundant, perceptible, and accurately identified by nature app observers, but which are almost
entirely limited to the river valley and riparian zones, include Bryoria, Cladonia, Parmelia, Peltigera,
Ramalina, and Vulpicida. Their limited distribution likely excludes them as biomonitors for air quality, but
they may be useful in monitoring climatic or other ecological shifts in urban parklands. The gradient
exerting the strongest effect on lichen community composition in Edmonton may be climate, a hypothesis
we will test in a future study.
At the species level, trained novices were successful surveying for Xanthomendoza fallax and the
abundant but morphologically variable Phaeophyscia orbicularis. Xanthomendoza is an eye-catching
genus, and given the rarity of the two other species documented in Edmonton (X. fulva and X. hasseana),
focusing hypothetical surveys on X. fallax is a logistically and ecologically acceptable loss of taxonomic
precision. Our results align with Sivanesan et al. (2005), who found that high school students were able to
conduct highly repeatable, focused lichen surveys, both in blind comparisons between students and
between students and instructors.
Based on our analyses of repeatability, field identification of individual Physcia species proved
difficult. Preliminary observations suggest that P. aff. stellaris and P. adscendens have broader niches than
P. aipolia group and P. aff. dimidiata. Future work will explore the loss of information in surveying for
Physcia at the genus level.
Another genus commonly included on bioindicator lists but that may be too challenging for
novice surveyors in Edmonton is Candelaria. Candelaria showed low detectability (Table 3), low
perceptibility (mostly misidentified and only one confirmed observation within the apps), and the two
species present appear to have divergent niches, which means that misidentifications would obscure
ecological information their presence may contribute. Whereas C. pacifica is common in the river valley
parks, we made multiple collections of C. concolor s.l. on boulevard trees in the tablelands. More
collections are needed to test this niche hypothesis.
Some crustose species or genera may be good biomonitors, but not surprisingly they were under-
represented in both student-based systematic surveys and within nature-reporting apps. Future work will
explore the subset of sites surveyed by more experienced lichenologists for the responsiveness of crustose
lichens to ecological gradients as well as potential groupings that could be tested with novice surveyors in
Edmonton.
We sourced two EMAN indicator lichen lists, one for mixed hardwood forests and one for boreal
forests (Brodo & Craig undated; D. McCarthy, unpublished). In total, these lists recommended 45 species
for air quality monitoring in Canada. When comparing our Edmonton lichen list to the composite EMAN
indicator species list, there was an overlap of only 21 species. The other 24 species were not detected in
Edmonton. Of those 21 overlapping species, ten were restricted to river valley and ravine parks. Of the
remaining 11 species, two are rare in the tablelands (Evernia mesomorpha, Parmelia sulcata) and two are
59
difficult to detect (Candelaria concolor s.l., Xanthomendoza hasseana). These findings illustrate the
importance of grounding biomonitoring studies in a strong foundation of local knowledge.
CONCLUSION
This is one of the first detailed studies of urban lichens and allied fungi diversity in continental
North America. Survey methods complemented each other: nature-reporting apps and opportunistic surveys
contributed unique records from habitats frequented by the public, while systematic surveys provided data
on species’ distribution and relative abundance across the city (Fig. 1). Molecular barcode data allowed us
to confirm or correct some identifications, and illuminated taxa that require additional phylogenetic work
and sampling. These datasets also provide guidance on which species are candidate bioindicators, i.e., those
species or genera that are both broadly distributed and easily identifiable (Table 4). Future work will
explore the sensitivity of these target lichens to known ecological and contaminant gradients in Edmonton.
ANNOTATED LIST OF SPECIES AND KEYS
In a departure from convention, the following list is organized first into 14 morphological groups
that we considered amenable to use by novices. Species are listed alphabetically within those groups. A
spreadsheet is also provided so that the list may be searched or organized by the reader (Supplementary
Appendix 1).
Many collections were mixed and are thus cited under multiple species. We also include
observations of specimens not collected (indicated with “unvouchered observation”) when we deemed
them reliable, including visually verified reports from nature-reporting applications. Collections of some
species are limited because of rarity, demonstrated ability to identify with confidence in the field, and/or
reluctance to sample from a highly visible location within a city park. Unless indicated otherwise, all
collections cited were examined for this study by the authors. All spot test results reflect testing done on
Alberta material. TLC results without references indicate analyses of Alberta material, largely through
work by D. Thauvette and D. Haughland through the ABMI.
We indicate uncertainty in the application of a given name in one of two ways. We use “cf.” (Latin
indicating to confer or to compare) where further study is required, and “aff.” (Latin ex affiniatis) where
taxonomic work involving Alberta material is ongoing and evidence to date suggests that the species in
question is not identical to the type specimen and represents a currently undescribed species.
Dichotomous keys are provided with caveats. The keys include only species currently confirmed
for Edmonton; including all possible additional or easily confused species would be beyond the scope of
this paper. As our knowledge of Edmonton’s flora is incomplete, users are cautioned to use the keys herein
as a starting point, and to then check their specimen against the included detailed species descriptions. If
any traits do not fit, they should consult the key literature listed for that group; their collection may
represent a new species for Edmonton. The first author welcomes all additional records and feedback.
Edmonton distribution maps for all species are provided in alphabetic order Supplementary Appendix 3
after the literature cited section of this contribution.
SUMMARY OF NOTATIONS AND ABBREVIATIONS
* = New to Alberta; ** = New to North America; *** = New to science
† = non-lichenized fungi
aff. = affinity
cf. = confer
s.l. = sensu lato, in the broad sense
s.s. = sensu stricto, in the strict sense
UoA-CC-# = University of Alberta Class Collections (students did not create individually numbered
collections; instead, all collections made during the biomonitoring surveys were contributed to and
renumbered as part of a UofA-CC collection)
60
MORPHOGROUP KEY
1a. Crustose lichens, thallus immersed in substrate or forming a thin episubstratic to areolate crust that
cannot be separated from the substrate intact .................................................................................................. 2
1b. Not as above, thallus foliose (leaf-like, with distinct upper and lower surface) or fruticose (three-
dimensional, hair-like, shrubby, cupped, or like a tiny dressmaking pin) ...................................................... 8
2a. Apothecial rim and disk dark grey, black, or brown.................................................................................. 3
2b. Apothecia at least in part orange, yellow, or white, or apothecia lacking ................................................. 5
3a. Apothecial rim easily visible and contrasting in color with the disk (lecanorine), the rim typically grey-
brown to dark brown with a dark disk ...................... Group 1: Brown-rimmed lecanorine crustose lichens
3b. Apothecia often black, either both rim and disk concolorous (lecideine) or with rim apparently lacking
(biatorine) ....................................................................................................................................................... 4
4a. Apothecia circular in outline .................................. Group 2: Lecideine and biatorine crustose lichens
4b. At least some apothecia elongate, script-like .................................................. Group 3: Lirellate lichens
5a. Apothecia with yellow disk and rim, K- ........................... Group 4: Yellow lecanorine crustose lichens
5b. Apothecia various but not with yellow disk and rim, K-, K+yellow or K+purple .................................... 6
6a. Apothecia with orange disk and/or rim, orange tissues K+purple at least in part .......................................
............................................................................................... Group 5: Orange lecanorine crustose lichens
6b. Apothecia with white rim (typically with a contrasting disk color, often with a white thallus), or
apothecia lacking, K- or K+yellow .................................................................................................................. 7
7a. Apothecia with white rim and contrasting disk ..................................................... Group 6: Lecanora s.l.
7b. Thallus sorediate or granular, typically lacking apothecia .........................................................................
.......................................................................................... Group 7: Sorediate or granular crustose lichens
8a. Thallus leaf-like (foliose), two-dimensional, often with distinct upper and lower surfaces ...................... 9
8b. Thallus three-dimensional (fruticose) at least in part, may have a crustose or squamulose primary
thallus, generally cortex similar throughout .................................................................................................. 13
9a. Leaf-like lichens of various colors, lobes typically ≤1cm wide, green algal photobiont only, becoming
greener when moistened ................................................................................................................................ 10
9b. Thallus with cyanobacterial photobiont at least in part, either in wart-like cephalodia or as dominant
photobiont in thallus, becoming darker (grey or black) when moistened; size varied................................... 12
10a. Leaf-like thallus orange, bright yellow or usnic pale yellow, with the latter reacting KC+ oily yellow ...
.................................................................................................. Group 8: Orange and yellow foliose lichens
10b. Leaf-like thallus predominantly white, grey, brown or green, KC-....................................................... 11
11a. Leaf-like lichens that are predominantly white or grey ......... Group 9: White and grey foliose lichens
11b. Leaf-like lichens that are predominantly brown to green . Group 10: Brown and green foliose lichens
12a. Lobes <0.5 cm wide, on soil or rock, black and gelatinous when wet ... Group 11: Small cyanolichens
12b. Lobes ≥ 0.5 cm wide and typically much larger, apothecia when present forming saddle-like structures
at the lobe tips, lower surface with rhizines and/or veins ................................................ Group 12: Peltigera
13a. Thallus ≤ 2 mm tall, minute fruiting bodies that resemble dressmaker pins .............................................
........................................................................................................... Group 13: Calicioid lichens and fungi
13b. Thallus >1 cm long/tall, fruiting bodies vary from globose terminal proliferations to disks ................ 14
14a. Thallus in the shape of cups, wands, shrubs, or clubs, mostly hollow, often found growing from a
squamulose or crustose primary thallus, common on soil, downed wood, and the bases of trees .....................
......................................................................................................................................... Group 14: Cladonia
14b. Thallus form hair-like, or coral-like thalli, mostly with solid interior, typically with a single
attachment point, either erect or pendulous, commonly epiphytic .... Group 15: Epiphytic fruticose lichens
61
GROUP 1: BROWN-RIMMED LECANORINE CRUSTOSE LICHENS
Four species. Key literature: McCune 2017a, 2017b; Sheard 2010; Sheard 2018; Sheard & May
1997. For confident identification, examination of spores under oil immersion (1000x) is necessary.
Rinodina albertana may also be present, but additional collections are required for confirmation.
1a. Spores constricted at septum, forming a figure-eight in outline; hypothecium red-brown ..........................
......................................................................................................................................Amandinea dakotensis
1b. Spores not constricted at septum; hypothecium clear to pale brown ......................................................... 2
2a. Immature spores with septal and apical wall thickenings, becoming thin walled with maturity
(Physconia-type spores); spores often slightly curved or “bean-shaped” ............................... Rinodina pyrina
2b. Spores with apical walls remaining thick through maturity ...................................................................... 3
3a. Spores not swelling at septum in K (Physcia-type spores) .................................................. Rinodina freyi
3b. Spores swollen at septum, becoming more pronounced in K (Dirinaria-type spores) ................................
......................................................................................................................................... Rinodina metaboliza
*Amandinea dakotensis (H. Magn.) P. May & Sheard FIGURE 12.
Apparently rare river valley epiphyte. New to Alberta, a single collection of what was initially
thought to be Rinodina was made from the bark of a downed tree. The spores indicated otherwise; rather
than having smooth walls typically thickened at least in part, these spores were minutely ornamented,
uniformly thin-walled, and had a distinctly constricted septum, creating a figure-eight shape. Widely
distributed in the eastern interior of the United States into southern Ontario, Sheard and May (1997)
reported this species from a similar latitude in the neighboring province of Saskatchewan. A prior
collection may exist from Wagner Natural Area east of Edmonton (Derek Johnson, pers. comm.), however,
the collection could not be located.
Edmonton material: grey-brown low areoles with closely aggregated lecanorine apothecia.
Apothecia lecanorine, <0.5 mm diameter, epihymenium dark brown, hymenium hyaline, hypothecium red-
brown. Thalline exciple scurfy, grey-brown. Spores 10–13.5 × 7–8 µm, brown, uniseptate, ovate, and
constricted at the septum, minutely rugose/ornamented, ornamentation visible only at 1000x magnification,
8 per ascus. Chemistry: all spot tests negative, no secondary metabolites detected by TLC (Sheard and May
1997). Molecular support: none, no sequences in GenBank, no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Sir Wilfrid Laurier Park, 2019,
53.50834, -113.560926, on Populus branches on downed tree, D. Haughland 2019-121D & P. Williams
(hb. Haughland).
Rinodina freyi H. Magn. FIGURE 13 E-F.
Apparently rare tableland epiphyte. Edmonton material: copper-brown areoles to 0.75 mm in
diameter, with 1–3 aggregated lecanorine apothecia. Spores 14–17 × 7–8 µm, brown, uniseptate, Physcia-
type development (Sheard 2010) with apical walls remaining thick throughout development, creating an
hourglass shape within the spore, unornamented, 8 per ascus. Spores often exhibit a well-developed torus,
an electron-dense, pigmented band at the septum (Sheard 2010). Hypothecium hyaline, epihymenium
brown to red-brown. Chemistry: all spot tests negative, no secondary metabolites detected by TLC (Sheard
2010). Molecular support: none, one mtSSU sequence in GenBank, no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 56,
53.520506, -113.570687, 2019, on trunk of Tilia, D. Haughland & A. Hood s.n. [UoA-CC-66] (hb.
Haughland); Sherwood Park, Air Quality Monitoring Station, 53.532016, -113.321511, 2019, on trunk of
Fraxinus pennsylvanica, D. Thauvette & J. Birch s.n. [UoA-CC-105] (hb. Haughland).
Rinodina metaboliza Vain. FIGURE 13 C-D.
Apparently rare tableland epiphyte. Edmonton material: clustered apothecia with prominent, raised
grey-brown thalline margins. Spores 14–22 × 7–9 µm, brown, uniseptate, some spores lightly ornamented,
62
Figure 12. Amandinea dakotensis morphology and anatomy, Edmonton, Haughland 2019-121D. A,
Thallus dry. B, Thallus wet. C, Spores showing minute warty ornamentation, thick spore wall, and
constricted septum. D, Apothecial cross section in water mount illustrating brown hypothecium, and
thalline exciple.
with Dirinaria-type development (apical walls remaining thick throughout development, creating an
hourglass shape within the spore, Sheard 2010), 8 per ascus. At maturity the spore tips become mucronate
(with an elongated nipple-like projection) and the spores widen at the septum, the latter becoming more
pronounced with the addition of K. Chemistry: All spot tests negative, no secondary metabolites detected
by TLC (Sheard 2010). Molecular support: none, four ITS sequences in GenBank, no new sequences
generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, South Air Quality Monitoring
Station, 53.501914, -113.524177, 2019, on trunk of Fraxinus, D. Thauvette & J. Birch s.n. [UoA-CC-104]
(hb. Haughland); Sherwood Park, Air Quality Monitoring Station, 53.532016, -113.321511, 2019, on trunk
of Fraxinus pennsylvanica, D. Thauvette & J. Birch s.n. [UoA-CC-105] (hb. Haughland); Edmonton,
University of Alberta North Campus, 53.527244, -113.519258, 2021, on Pinus sylvestris, J. Singh & K.
Schafer s.n. [UoA-CC21-23].
Rinodina pyrina (Ach.) Arnold FIGURE 13 A-B.
Common tableland epiphyte. While this is overwhelmingly the most common Rinodina on open-
growing trees in Edmonton, collections should be critically examined to exclude rarer species. Edmonton
material: variable in morphology but commonalities include clustered apothecia with visible thalline rims
with relatively large-celled trebouxioid algae in cross-section, and a dark brown, epruinose disk. The
thalline exciple varied in color from brown to grey-brown. Apothecia characterized by the brown pigments
63
Figure 13. Rinodina species of Edmonton, thallus (dry) and spores in water mount. A-B, R. pyrina,
showing curved spores, UoA-CC-26. C-D, R. metaboliza, showing widening at septum, UoA-CC-105. E-F,
R. freyi, showing torus. E, UoA-CC-66, F, UoA-CC-105.
in the epihymenium, the hyaline hymenium and hypothecium, and brown, uniseptate spores 12–14 × 6–7
µm, 8 per ascus. Spores are Physconia-type (Sheard 2010), with thin spore walls and rounded locules at
maturity, typically with a visible torus, and often curved (“kidney bean”-shaped) at maturity, with no
swelling or constriction at the septum. Chemistry: All spot tests negative, no secondary metabolites
64
detected by TLC (Sheard 2010). Molecular support: none, >10 sequences in GenBank, no new sequences
generated.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 3, 53.440376, -113.484487, 2019, on trunk of Ulmus americana, D. Thauvette & J. Wasyliw s.n. [UoA-
CC-108] (hb. Haughland); Edmonton, Urban Monitoring Site 33, 53.479311, -113.413258, 2019, on trunk
of Tilia, D. Thauvette et al. s.n. [UoA-CC-46,81] (hb. Haughland); Edmonton, Urban Monitoring Site
107E, 53.548554, -113.617129, 2019, on trunk of Salix, D. Haughland s.n. [UoA-CC-97] (hb. Haughland);
Edmonton, Urban Monitoring Site 119, 53.602014, -113.458033, 2019, on trunk of Fraxinus
pennsylvanica, D. Royko & D. Fielder s.n. [UoA-CC-176] (hb. Haughland); Edmonton, Urban Monitoring
Site 121E, 53.561280, -113.609950, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland s.n. [UoA-
CC-10] (hb. Haughland); Edmonton, Urban Monitoring Site 52E, 53.494372, -113.597330, 2019, on trunk
of Ulmus americana, D. Haughland s.n. [UoA-CC-21] (hb. Haughland); Edmonton, Beverly Air Quality
Monitoring Station, 53.566860, -113.397464, 2019, on trunk of Ulmus americana, D. Thauvette & J. Birch
s.n. [UoA-CC-26] (hb. Haughland); Edmonton, South Air Quality Monitoring Station, 53.501946, -
113.5249, 2019, on trunk of Fraxinus, D. Thauvette & J. Birch s.n. [UoA-CC-132] (hb. Haughland).
GROUP 2: LECIDEINE AND BIATORINE CRUSTOSE LICHENS
Nine species. Key literature: Björk 2013; Ekman 1996; McCune 2017a, 2017b; Smith et al. 2009.
This group is under-collected, with a relatively high species discovery rate in examined material, and three
species new to Alberta. Another species reported for Edmonton that we could not confirm is Micarea
melaena (under Bacidia melaena, Rainbow Valley, Edmonton, 1961, G.W. Scotter 706 [CANL], reported
in BIOTICS [Government of Alberta 2020], not examined here). For confident identification, examination
of apothecia cross-sections and spores with a compound microscope is necessary.
1a. Apothecia biatorine, beige, pink, yellow, grey or dark-red to almost black, proper exciple often visible
and paler than disk ........................................................................................................................................... 2
1b. Apothecia lecideine, with a black disk and black proper exciple .............................................................. 5
2a. Growing on soil and mosses, sometimes on tree bases but not directly on bark; hypothecium orange-
brown in apothecia cross-sections; spores with 3–5 transverse septa and minute warty ornamentation on
perispore ........................................................................................................................ Bilimbia sabuletorum
2b. Epiphytic, growing directly on/in bark; hypothecium hyaline in apothecia cross-sections; spores with
≤3 transverse septa, unornamented .................................................................................................................. 3
3a. Apothecia beige to yellow; on a grey-blue, rough, verrucose thallus .......................... Lecanora symmicta
3b. Apothecia pink, piebald or darkening to brick-red or black; on a pale green thin to areolate episubstratic
thallus .............................................................................................................................................................. 4
4a. Spores simple .......................................................................................................... Lecidea erythrophaea
4b. Spores with 1-3 transverse septa ..................................................................................... Lecania naegelii
5a. Thallus dark-green and granular; spores curved to S-shaped, ≥3 septate, spiralling around each other
within the asci ........................................................................................................ Scoliciosporum umbrinum
5b. Thallus white to grey-green, varying from immersed to dust-like, verrucose, or placodioid; spores never
S-shaped .......................................................................................................................................................... 6
6a. Thallus white, immersed to dust-like; hypothecium black in apothecia cross-sections; spores with one
transverse, constricted septum and cells of unequal size ................................................. Arthonia patellulata
6b. Thallus grey and episubstratic at least in part; hypothecium hyaline or rust to orange-brown in
apothecia cross-sections; spores varying from simple to septate but cells of equal size and septa not
constricted ....................................................................................................................................................... 7
7a. Spores simple; hypothecium rust to orange-brown ............................................... Lecidella elaeochroma
7b. Spores transversely septate; hypothecium hyaline .................................................................................... 8
8a. Spores >8 per ascus, bean-shaped or slightly curved, ≤12 µm in length, with 1–3 transverse septa;
common ................................................................................................................. Arthrosporum populorum
8b. Spores 8 per ascus, narrowly bacilliform, ≥18 µm in length, with ≥3 transverse septa; rare ......................
........................................................................................................................................Bacidia circumspecta
65
Figure 14. Arthonia patellulata morphology and anatomy, Edmonton, UoA-CC-71. A, Apothecial cross
sections – top in water, bright light; middle section in water, polarized light; bottom section treated with KI.
B, Habit, on Populus tremuloides in parkland habitat. C, Spores in water mount, showing unequal cell size.
Arthonia patellulata Nyl. FIGURE 14.
Apparently rare river valley epiphyte. Edmonton material: thallus white and dust-like to immersed
and non-apparent, with scattered black, matte lecideine apothecia. Spores hyaline, uniseptate, narrowly
obovate, septum constricted, cells of unequal size, unornamented, 10–12 × 4–5 µm, 8 per ascus. Hymenium
hyaline, ~65 µm thick, epihymenium faintly black, paraphyses without brown caps, hypothecium black,
proper exciple black, poorly developed. Asci with well-developed torus. Apothecia slightly convex.
Photobiont trebouxioid. Hymenium K+brownish, K/I+blue. Chemistry: all thallus spot tests negative, no
lichen substances known (Björk 2013). Molecular data: 3 eDNA nrITS sequences in GenBank, 1 new
sequence generated (isolate DLH39 from UoA-CC-71), however BLAST indicated it is most similar to
GenBank-accessioned Candelariella vitellina, perhaps due to a processing or sampling error. No further
analyses conducted.
Specimen examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 23E,
53.478452, -113.620128, 2019, on trunk of Populus, D. Haughland s.n. [UoA-CC-71] (hb. Haughland).
66
Figure 15. Arthrosporum populorum morphology and anatomy, Edmonton. A, Habit, on Fraxinus
pennsylvanica, UoA-CC-1. B, Habit, on Ulmus americana growing in a grassy boulevard, UoA-CC-120. C,
Asci and spores in water mount, UoA-CC-1. D, Spores in water mount, UoA-CC-93.
Arthrosporum populorum A. Massal.
(≡ Toninia populorum (A. Massal.) Kistenich, Timdal, Bendiksby & S. Ekman; Kistenich et al. 2018)
FIGURE 15.
Common tableland and river valley and ravine system epiphyte on a diversity of deciduous tree
species. Björk (2013) hypothesized that Alberta material is not A. populorum s.s. as the apothecia and
spores are smaller than reported elsewhere, and tentatively called it A. “nanum Björk ined.” Edmonton
material: thallus indistinct to grey placodioid, with black lecideine apothecia, diameter to 0.54 mm, proper
margin often persistent, well-developed. Spores 9–12 × 3–5 µm, 1–3 transverse septate, rounded ends,
hyaline, slightly curved, >8 per ascus. Photobiont trebouxioid. Hyaline hymenium and hypothecium,
epithecium black, proper exciple black externally and pale rusty brown internally. Paraphyses free in water,
slightly capitate, septate. Chemistry: all thallus spot tests negative, no lichen substances known (Björk
2013). Molecular support: Kistenich et al. (2018) show this species in a well-supported clade of Toninia
species, no new sequences generated.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 17, 53.466685, -113.571654, 2019, on trunk of Populus, D. Haughland & L. Hjartarson s.n. [UoA-
CC-98] (hb. Haughland); Edmonton, Urban Monitoring Site 36, 53.494272, -113.620171, 2019, on trunk of
Fraxinus pennsylvanica, D. Haughland & L. Hjartarson s.n. [UoA-CC-1] (hb. Haughland); Edmonton,
Urban Monitoring Site 45E, 53.480176, -113.429404, 2019, on trunk of Populus hybrid, S. Toni & M.
Lewis s.n. [UoA-CC-93] (hb. Haughland); Edmonton, Urban Monitoring Site 86, 53.561777, -113.547997,
2019, on trunk of Ulmus americana, D. Haughland & A. Hood s.n. [UoA-CC-120] (hb. Haughland);
Edmonton, Urban Monitoring Site 127, 53.6111, -113.4608, 2019, on trunk of Ulmus, D. Haughland s.n.
[UoA-CC-41] (hb. Haughland); Edmonton, Urban Monitoring Site 149E, 53.6111, -113.4608, 2019, on
67
Figure 16. Bacidia circumspecta morphology and anatomy, Edmonton. A, Thallus, wet, on Populus
balsamifera, Haughland 2021-1. B, Asci treated with K followed by Lugol’s I, mature (left) and immature
(right), UoA-CC-84. C, Apothecial cross-section under white light, UoA-CC-84. D, Spores in water mount,
UoA-CC-84.
trunk of Salix cf. pentandra, D. Haughland & L. Hjartarson s.n. [UoA-CC-100] (hb. Haughland);
Ardrossan, Air Quality Monitoring Station, 53.554824, -113.143457, 2019, on trunk of Populus cf.
balsamifera, D. Thauvette & J. Birch s.n. [UoA-CC-69] (hb. Haughland).
*Bacidia circumspecta (Nyl. ex Vain.) Malme FIGURE 16.
Apparently rare river valley and parkland epiphyte. New to Alberta, the nearest prairie province
record is from Prince Albert National Park, Saskatchewan (Ekman 1996). Edmonton material:
discontinuous grey to grey-green to brown areolate thallus. Apothecia lecideine, to 1 mm diameter, cup-
shaped when immature, becoming flat to slightly convex, disc black and concolorous with proper exciple,
epruinose. Excipular cells thick-walled, wider towards rim, clearly differentiated from paraphyses.
Paraphyses black capitate, to 4 µm wide at tip, simple to sparsely branched at very tip. Spores 18–22 × 3
µm, straight, bacilliform (sensu Ekman 1996), hyaline, unornamented, mostly 3–4 transverse septa,
occasionally up to 6 septa present, 8 per ascus. No polarizing crystals in apothecia cross-sections. Co-
occurring with Phaeophyscia kairamoi in both collections. Epihymenium and proper exciple N+reddish-
purple, K-, C-, hypothecium hyaline, N+ yellowish, K-, C-. Chemistry: all thallus spot tests negative, no
secondary metabolites detected (Ekman 1996). Molecular support: a single ITS sequence (isolate DLH35
from UoA-CC-84), is most similar to Bacidia circumspecta GenBank Accessions AF282124 (Sweden, 97%
percent identity, 12 positions different, 479 bp overlap, Ekman 2001) and MH539764 (Russia, 96% percent
identity, 21 positions different, 549 bp overlap, Gerasimova et al. 2018). These differences are smaller than
the difference between the two previously accessioned sequences (23 positions different, 476 bp overlap).
68
Figure 17. Bilimbia sabuletorum morphology and anatomy, Edmonton, Haughland 2020-61. A, Thallus
growing on mineral soil and moss, wet. B, Apothecial cross-section showing orange-brown hypothecium.
C, Asci and spores after treatment with K. D, Spores in water mount.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 47,
53.505866, -113.553095, 2019, on trunk of Populus balsamifera, D. Haughland s.n. [UoA-CC-86] (hb.
Haughland); Edmonton, Urban Monitoring Site 47, 53.505866, -113.553095, 2019, on trunk of Populus
balsamifera, D. Haughland 2021-1(hb. Haughland); Edmonton, Urban Monitoring Site 163E, 53.608013, -
113.590864, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n. [UoA-CC-84] (hb.
Haughland).
Bilimbia sabuletorum (Schreber) Arnold
(≡ Mycobilimbia sabuletorum (Schreber) Hafellner) FIGURE 17.
Apparently rare river valley terricole. Edmonton material: thallus thin, pale grey-green, granular,
with abundant black-red brown to grey biatorine apothecia. Epithecium with fine, polarizing crystals,
hymenium hyaline to pale yellow, 100–150 µm thick, hypothecium orange-brown. Spores fusiform,
hyaline, perispore with minute warty ornamentation that can be difficult to see, 23–30 × 5–8 µm, with 3–5
transverse septa, swelling slightly in K, 8 per ascus. Chemistry: all thallus spot tests negative, no lichen
substances known (Björk 2013). Molecular support: genus and to a lesser extent species supported in
Kistenich et al. (2018), no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Hawrelak Trail off-leash park,
53.517853, -113.54035, 2020, on exposed mineral soil and moss, D. Haughland 2020-61 & K. Tichkowski
(hb. Haughland).
69
Figure 18. Lecania naegelii morphology and anatomy, Edmonton, UoA-CC-49. A, Thallus, on Populus
tremuloides. B, Apothecial cross-section under white (top) and polarized light (bottom). C, Spores in water
mount. D, Ascus treated with K followed by Lugol’s I.
*Lecania naegelii (Hepp) Diederich & van den Boom FIGURE 18.
Apparently rare river valley epiphyte. This collection fits within the documented niche of this
species: nutrient-rich, smooth-barked trees (van den Boom & Ryan 2004). While listed as unrankable on
Alberta’s List of Elements (rank SU, Government of Alberta 2017), we could not locate any previous
records so here we report it as new to Alberta. Edmonton material: thin whitish thallus with convex pale
green areoles. Apothecia plane, <0.5 mm in diameter, with a pink partially blackened disk and paler
contrasting rim. In cross-section, no polarizing crystals were found except near the algal layer at the base
and point of attachment; the rim was composed of hyaline, radiating thick-walled hyphae; the epithecium
was largely hyaline with dispersed black pigments on the slightly swollen tips of the septate, simple
paraphyses. Hymenium and hypothecium hyaline. Asci biatorine-type, staining K/I+blue, with an
elongated, non-amyloid axial mass in a darkly amyloid tholus. Spores 1-3 transverse septate, hyaline,
straight to slightly curved or bean-shaped, 12–17 × 4–5 µm, 8 per ascus, with rounded ends. Photobiont
trebouxioid, restricted to the base of the apothecia. Chemistry: all thallus spot tests negative, no secondary
metabolites detected (van den Boom & Ryan 2004). Molecular support: A single ITS sequence (isolate
DLH14 from UoA-CC-49) has >97% percent identity with four accessioned Lecania naegelii GenBank
sequences (Accession AM292691, 12 positions different, 496 bp overlap; KT695396 & KT695323,
Canada, 15 positions different, 547 bp overlap; FR799198, United Kingdom, 15 positions different, 521 bp
overlap).
70
Figure 19. Lecanora symmicta s.l. morphology and anatomy, Edmonton. A, Thallus on dead Alnus,
Haughland 2020-15A. B, Apothecial cross-section in polarized light showing fine crystals that dissolve in
K, Haughland 2020-15A. C, Ascus, UoA-CC21-18. D, Apothecial cross-section in white light, Haughland
2020-15A.
Specimen examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 23E,
53.470689, -113.620142, 2019, on trunk of Populus tremuloides, D. Haughland s.n. [UoA-CC-49] (hb.
Haughland).
Lecanora symmicta (Ach.) Ach. s.l. FIGURE 19.
Apparently rare river valley and tableland epiphyte, likely present in parkland. Edmonton
material: thallus pale blue-grey, episubstratic in part with small, rough verrucae that give rise to apothecia
and/or dissolve into coarse granules. Apothecia biatorine, beige to yellow, convex and crowded, becoming
“molten” and fused, no algae found in margin, proper exciple thin. Spores non-septate to rarely 1-septate,
hyaline, unornamented, ellipsoidal, 8–12 × 4–5 µm. Polarizing crystals present in epihymenium and
subhymenium that dissolve in K. Hymenium K/I+ fleeting blue. Chemistry: all thallus spot tests negative,
usnic acid, zeorin, arthothelin, theophanic acid, 4,5-dichloronorlichexanthone (trace), norlichexanthone
(trace) by TLC (Ryan et al. 2004). Molecular data: no new sequences generated. Given our material is
atypical in some respects (occasional septate spores, granules) and other species have been recently split
from this species (Pérez-Ortega & Kantvilas 2018), it is a priority for future molecular work.
Specimens examined. – CANADA. ALBERTA: Edmonton, Whitemud Ravine, 53.491661, -
113.55914, 2020, on bark of dead Alnus snag, D. Haughland 2020-15A & P. Williams (hb. Haughland);
Edmonton, University of Alberta North Campus, 53.525395, -113.525717, 2021, on Pinus mugo, J. Singh
71
Figure 20. Lecidea erythrophaea morphology and anatomy, Edmonton, UoA-CC-67. A, Thallus on large-
bole Populus balsamifera. B, Apothecial cross-section under white (bottom) and polarized light (top). C,
Asci and spores in water mount. D, Ascus treated with K followed by Lugol’s I.
& K. Schafer s.n. [UoA-CC21-18] (hb. Haughland); Edmonton, University of Alberta North Campus,
53.5254522, -113.526155, 2021, on Pinus mugo, K. Schafer & J. Singh s.n. [UoA-CC21-29] (hb.
Haughland); Edmonton, MacKenzie Ravine, 1976, on rotting log, D.C. Lindsay s.n. (PMAE- B77.24.41).
*Lecidea erythrophaea Flörke ex Sommerf. FIGURE 20.
Apparently rare river valley epiphyte. New to Alberta, we detected a single specimen from the
North Saskatchewan River Valley on a large diameter Populus balsamifera. The closest known collection
is from Wells Gray Provincial Park, British Columbia, from a swamp forest (C. Björk 25583, NY [n.v.],
CNALH 2020). Edmonton material: apothecial disk brick-red to dark brown, thallus pale greenish-grey,
smooth, thin. Spores non-septate, 8–9 × 2.5–3 µm, hyaline, unornamented, narrowly ellipsoid, 8 per ascus.
Epihymenium tan brown, hymenium and subhymenium hyaline. Chemistry: all thallus spot tests negative,
no secondary metabolites detected by TLC (Hertel & Printzen 2004). Molecular data: no sequences in
GenBank, no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 57x,
53.521192, -113.548532, 2019, on trunk of Populus balsamifera, D. Haughland & A. Hood s.n. [UoA-CC-
67] (hb. Haughland).
72
Figure 21. Lecidella elaeochroma morphology and anatomy, Edmonton, A-B, D. Haughland 2020-43, D-
C, UoA-CC-125. A-B, Habit, on downed Betula. C, Apothecial cross-section under white (left) and
polarized light (right). D, Asci and spores in water mount, inset treated with K followed by Lugol’s I.
Lecidella elaeochroma (Ach.) M. Choisy FIGURE 21.
Occasional river valley and ravine system and parkland epiphyte. Edmonton material: thallus
thinly verrucose to cracked, grey-green, with black prothallus in some collections, with black lecideine
apothecia with distinct exciple when young, becoming convex with age, to 1 mm diameter. Spores simple,
10–14 × 5–8 µm, broadly ellipsoid to lemon-shaped, hyaline, unornamented, 8 per ascus. In apothecia
cross-sections the epihymenium and proper exciple are blue-black, the exciple cells are hyaline to almost
violet and not carbonized, the hypothecium is rusty/orange-brown, and the hymenium hyaline. Apothecium
interspersed with crystals in hypothecium and thinly in epihymenium. Asci with axial body and K/I+blue
thollus. Chemistry: thallus K+ yellow, PD-, C-, KC-, complex of xanthones by TLC (Knoph and Leuckert
2004). Molecular support: we originally identified this material as Lecidella euphorea (Flörke) Hertel, but
redetermined it based on our analyses of a single ITS sequence (isolate DLH11 from Haughland 2020-43).
Previous authors have suggested that molecular data or high-performance liquid chromatography is
required to definitively differentiate these two species (e.g., McCune 2017b). The L. elaeochroma clade is
polyphyletic, and our western North American sequences fall on a well-supported branch separate from but
close to a clade with L. “elaeochroma 5” from Europe (Zhao et al. 2015; Fig. 6 herein).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 56, 53.520506, -113.570687, 2019, on trunk of Tilia, D. Haughland & A. Hood s.n. [UoA-CC-66] (hb.
Haughland); Edmonton, Urban Monitoring Site 148E, 53.587522, -113.640605, 2019, on trunk of Populus
balsamifera, D. Haughland & L. Hjartarson s.n. [UoA-CC-125] (hb. Haughland); Edmonton, Urban
Monitoring Site 57X, 53.521192, -113.548532, 2019, on trunk of Populus balsamifera, D. Haughland & A.
73
Figure 22. Scoliosporum umbrinum morphology and anatomy, Edmonton, UoA-CC21-18. A, Thallus, on
Pinus mugo bark. B, Apothecial cross-section under white light with Leica optical shading, showing spores
spiralling in ascus. C, Asci and branched paraphyses in water mount. D, Spore in water mount.
74
Hood s.n. [UoA-CC-67] (hb. Haughland); Edmonton, Patricia Ravine, 53.503105, -113.592863, 2020, on
bark of recently downed Betula papyrifera, D. Haughland 2020-43 & A. Hood (hb. Haughland);
Edmonton, University of Alberta North Campus, 53.527244, -113.519258, 2021, on Pinus sylvestris, K.
Schafer & J. Singh s.n. [UoA-CC21-23] (hb. Haughland).
Scoliciosporum umbrinum (Ach.) Arnold FIGURE 22.
Apparently rare tableland epiphyte. Edmonton material: thallus of dark-green, matte granules,
lacking soredia, isidia or prothallus. Apothecia black, shiny, to 0.25 mm wide, with proper exciple (evident
when young, becoming largely excluded), apothecia becoming slightly convex with age. Branching, septate
paraphyses in gelatinous matrix, not swollen at tips, 3 µm wide, also forming proper exciple. Asci
Lecanora-type. Epithecium grey to blue-green, hymenium and hypothecium hyaline. Spores hyaline,
unornamented, fusiform, 23–30 × 3–3.5 µm, 3–4 septate, mostly curved to S-shaped, spiralling in asci,
apparently 8 per ascus but hard to be definitive. Apothecial tissues in wet mount largely lacking polarizing
crystals (a few tiny crystals on surface), K-, asci KI+blue, hypothecium and paraphyses KI-. Chemistry: all
spot tests on the thallus negative, no lichen substances known (Björk 2013). Molecular support: good
genus-level and limited species-level support (Fryday et al. 2020), no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, University of Alberta North Campus,
53.5253952, -113.525717, 2021, on trunk of Pinus mugo, J. Singh & K. Schafer s.n. [UoA-CC21-18] (hb.
Haughland); Edmonton, grassy park next to Saskatchewan Drive bordering river valley, 53.513211, -
113.538619, 2021, on Picea twigs, D. Haughland 2021-30 (hb. Haughland).
GROUP 3: LIRELLATE LICHENS
One species. Key literature: Ertz & Egea 2007; Torrente & Egea 1989.
Alyxoria varia (Pers.) Ertz & Tehler
(≡ Opegrapha varia Pers.) FIGURE 23.
Occasional river valley and ravine system epiphyte. Rediscovered for Alberta; see the results
section for more information on prior reports. Extensive colonies were found on mature Populus
balsamifera in the riparian zone of creeks and the North Saskatchewan River. Edmonton material: lirellae
on a farinose crust, photobiont appearing trebouxioud, individual algal cells to 15–20 µm in diameter.
Lirellae rounded, irregular or elongate, sessile, superficial, and slightly constricted at base, 0.2–1 mm long
and 0.16–0.3 mm wide. Lirellae lacking thalline exciple, proper exciple shiny, epruinose and strongly
carbonaceous so that anatomy obscured in section, curving over an open hymenium and appearing closed
beneath the subhymenium. The hymenium varies from epruinose to lightly greenish-yellow pruinose.
Spores narrowly obovate, 8 per ascus, 20–25 × 6–8 µm, mostly with 5 transverse septa and an enlarged
middle cell, cell walls only slightly thickened at septa. Spores hyaline with dark walls, halonate, at maturity
darkening and becoming warty. Apothecial tissues were almost exclusively K/I- except very minimal
K/I+blue reactions in limited areas of the hymenium in a single section. Chemistry: all thallus spot tests
negative, no secondary metabolites detected by TLC (Ertz & Egea 2007). Molecular support: a complex of
species in need of revision, sequences analysed to date forming a distinct clade within the genus (Ertz et al.
2020), no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.501505, -
113.601141, 2019, on bark of mature Populus balsamifera in riparian zone, D. Haughland et al. 2019-116
(hb. Haughland); Edmonton, Patricia Ravine, 53.504611, -113.593583, 2020, on trunk of live >75 cm DBH
Populus balsamifera along trail, D. Haughland 2020-54 & A. Hood (hb. Haughland); Edmonton, Rat
Creek, Kinnaird Ravine, 53.5582, -113.453925, 2020, on trunk of live Populus balsamifera along trail, D.
Haughland 2020-105 & P. Williams (hb. Haughland).
75
Figure 23. Alyxoria varia morphology and anatomy, Edmonton, Haughland 2019-116. A-B, Habit, on
Populus tremuloides in riparian zone. C, Green algae in farinose thallus. D, Asci and spores in water
mount. E, Cross-section of apothecium and substrate.
76
GROUP 4. YELLOW LECANORINE CRUSTOSE LICHENS
Four species. Key literature: Brodo 2016; McCune 2017a, 2017b; Westberg 2004, 2005, 2007a,
2007b; Westberg et al. 2011. Additional species from this group that may be present in Edmonton include
Candelariella xanthostigma, but existing collections are too small for confident determination. Preliminary
identification is possible using morphology and substrate alone; confident identification requires
examination of the spores. K is helpful in excluding co-occurring Group 5 Caloplaca s.l. (K+ purple in
Caloplaca s.l. vs. K- to K+ reddish in Candelariella).
1a. Growing on rock or concrete; 8 spores per ascus ..................................................... Candelariella aurella
1b. Growing on bark or wood or other anthropogenic substrates; 8 or more spores per ascus ....................... 2
2a. Thalllus grey, may appear minutely tomentose; proper exciple grey to yellow; 8 spores per ascus
................................................................................................................................. Candelariella antennaria
2b. Thallus yellow, never tomentose; proper exciple either yellow or not visible; 12 or more spores per
ascus ................................................................................................................................................................ 3
3a. Thallus of small, flat, dispersed areoles ≤0.1 mm wide; proper exciple not visible ....................................
.................................................................................................................................... Candelariella cf. lutella
3b. Thallus larger, becoming subsquamulose, to 0.5 mm wide, often aggregated; proper exciple typically
visible ...................................................................................................................... Candelariella cf. vitellina
*Candelariella antennaria Räsänen FIGURE 24 A-C.
Occasional tableland epiphyte. This species is reported as new to Alberta. We checked PMAE
under the names Candelariella aurella (Hoffm.) Zahlbr. and C. deflexa (Nyl.) Zahlbr. for specimens with
grey thalli but found none. This species is reported to have a wide ecological amplitude; alternatively, it
may represent a complex of similar species (Westberg & Sohrabi 2012). In the neighboring provinces of
British Columbia and Saskatchewan, it has been found on rock, soil, Populus snags, Artemisia, Ulmus, and
Krascheninnikovia (CNALH 2020, Freebury 2014), and the senior author observed it on Populus along the
South Saskatchewan River, east of Saskatoon (D. Haughland, 2020, unvouchered observation). In other
parts of its range including Russia, Kazakhstan, and Nepal, it grows on a wide variety of deciduous trees
and shrubs (Yakovchenko et al. 2012). Edmonton material: epiphytic on a variety of deciduous trees,
thallus pale to dark grey to green-grey, indistinct, thin, or thick and amorphous. Apothecia scattered to
crowded, 0.1–0.3 mm diameter, disk yellow, flat to somewhat convex, yellow or grey thalline margin
(smooth or appearing slightly tomentose), proper margin indistinct. Spores hyaline, simple to 1-septate,
narrowly ellipsoid, 11–17 × 5–7 µm, 8 per ascus. Chemistry: all spot tests negative, calycin, pulvinic acid,
pulvic acid lactone and vulpinic acid in yellow parts by HPLC (Westberg 2007b). Molecular data: limited
species-level support as a sister clade to C. aggregata M. Westb. (Liu et al. 2019) but more analyses
needed, no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 40,
53.493796, -113.504715, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-
CC-16] (hb. Haughland); Edmonton, Urban Monitoring Site 62, 53.520757, -113.432138, 2019, on trunk of
Ulmus americana, S. Toni & A. Hood s.n. [UoA-CC-28] (hb. Haughland); Edmonton, Urban Monitoring
Site 135, 53.642107, -113.502186, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n.
[UoA-CC-30] (hb. Haughland); Edmonton, Urban Monitoring Site 106E, 53.546421, -113.641504, 2019,
on trunk of Ulmus americana, D. Haughland s.n. [UoA-CC-103] (hb. Haughland); Edmonton, Urban
Monitoring Site 150E, 53.590394, -113.588685, 2019, trunk of Ulmus americana, D. Haughland & L.
Hjartarson s.n. [UoA-CC-27] (hb. Haughland); Edmonton, Sir Wilfrid Laurier Park, 53.50834, -
113.560926, 2019, on bark of large downed Populus, D. Haughland 2019-122C & P. Williams (hb.
Haughland); Edmonton, University of Alberta North Campus, 53.527511, -113.519763, 2021, on Aesculus
glabra, K. Schafer & J. Singh s.n. [UoA-CC21-46] (hb. Haughland).
77
Figure 24. Yellow lecanorine crustose Candelariella of Edmonton. A-C, C. antennaria, new to Alberta. A,
Specimen with well-developed thallus, UoA-CC-103. B, Typical specimen with thallus limited to areoles
bearing apothecia, Haughland 2019-122C. C, Asci, paraphyses and spores in water mount, UoA-CC-103.
D, C. aurella on sidewalk, Spribille https://www.inaturalist.org/observations/44151398. E, C. cf. vitellina
on Fraxinus pennsylvanica, UoA-CC-113. F, C. cf. lutella on bark of Populus balsamifera, UoA-CC-23.
78
Candelariella aurella (Hoffm.) Zahlbr. FIGURE 24 D.
Anthropogenic saxicole. Thallus of yellow to orange-yellow areoles with abundant lecanorine
apothecia. Spores hyaline, narrowly ellipsoidal, simple to 1-septate, unornamented, 13–16 × 6–8 µm, 8 per
ascus. Chemistry: thallus K- or KC+ reddish, C-, PD-, UV-, calycin, pulvinic acid, pulvinic dilactone and
vulpinic acid by TLC (Westberg 2004). Molecular data: species well-supported phylogenetically (Liu et al.
2019, Westberg & Arup 2011) no new sequences generated.
Specimen cited. – CANADA. ALBERTA: Edmonton, Strathcona, 9736 90 Ave. NW, 53.525645,
-113.481477, 2018, on sidewalk concrete, T. Spribille 42799 (hb. Spribille: iNaturalist record
https://www.inaturalist.org/observations/44151398).
Candelariella cf. lutella (Vain.) Räsänen FIGURE 24 F.
Occasional river valley, parkland, and tableland epiphyte. Edmonton material: greenish yellow to
yellow small, flat areoles 0.05–0.1 mm wide, forming scattered or crowded patches 0.5–3.5 mm wide.
Apothecia 0.1–0.5 mm diameter, disk and thalline exciple colored like thallus, proper exciple indistinct,
thalline exciple often beaded. Spores 7–12 × 4–5 µm, simple, asci with 24–32 spores. Similar to C. cf.
vitellina but this species’ areoles are smaller, more dispersed, and typically do not form overlapping
aggregates, and the proper exciple usually is not visible. Chemistry: thallus K- or KC+ reddish, C-, PD-,
UV-, secondary metabolites not investigated (Westberg 2004). Molecular support: in our phylogeny a
single ITS sequence (isolate DLH39 from Haughland 2019-121A) does not group with C. lutella sequences
from Westberg (Fig. 2 herein). Instead it forms a highly supported basal branch to four identical sequences
determined as C. efflorescens (Westberg et al. 2007) and C. cf. vitellina (Fig. 2 herein). Additional work is
required to resolve the taxonomy of epiphytic Candelariella in Edmonton.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 28,
53.479694, -113.549752, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n.
[UoA-CC-52] (hb. Haughland); Edmonton, Urban Monitoring Site 40, 53.493790, -113.504979, 2019, on
trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-CC-8] (hb. Haughland); Edmonton,
Urban Monitoring Site 47, 53.505933, -113.552966, 2019, on trunk of Populus balsamifera, D. Haughland
s.n. [UoA -CC-23] (hb. Haughland); Edmonton, Urban Monitoring Site, 56, 53.520506, -113.570687, 2019,
on trunk of Tilia, D. Haughland & A. Hood s.n. [UoA-CC-66] (hb. Haughland); Edmonton, Urban
Monitoring Site 62, 53.520757, -113.432138, 2019, on trunk of Ulmus americana, S. Toni & A. Hood s.n.
[UoA-CC-28] (hb. Haughland); Edmonton, Urban Monitoring Site 150E, 53.590394, -113.588685, 2019,
on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n. [UoA-CC-27] (hb. Haughland);
Edmonton, Sir Wilfrid Laurier Park, 53.50834, -113.560926, 2019, on Populus twigs on downed tree, D.
Haughland 2019-121A & P. Williams (hb. Haughland).
Candelariella cf. vitellina (Hoffm.) Müll. Arg. FIGURE 24 E.
Common river valley, parkland, and tableland epiphyte. This is the most common epiphytic
Candelariella in Edmonton across a diversity of deciduous trees on boulevards and in river valley parks. In
other regions this species is more commonly found on non-calcareous rock (Westberg 2007a, but see Björk
2013), suggesting molecular work would be beneficial. Edmonton material: thallus of bright yellow
abundant granules or areoles, esorediate, varying from rounded and granular to subsquamulose, slightly
flattened and irregularly incised, often crowded and overlapping, forming pulvinate clusters. Apothecia
usually with the thallus but may be disjunct in space; apothecia bright yellow with similar colored thalline
exciple and distinct proper exciple. Spores 8–12 × 3–5 µm, simple or with a thin septum, 12–24 per ascus.
Chemistry: K- or KC+ reddish, C-, PD-, UV-, calycin, pulvinic acid, pulvinic dilactone and vulpinic acid in
yellow parts by TLC (Westberg 2004). Molecular support: three ITS sequences are identical to the single
published sequence of C. efflorescens (Westberg et al. 2007; Fig. 3 herein). Additional work is required to
resolve the taxonomy of epiphytic Candelariella in Edmonton.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 9, 53.453708, -113.527608, 2019, on trunk of Fraxinus, D. Haughland s.n. [UoA-CC-144] (hb.
Haughland); Edmonton, Urban Monitoring Site 45, 53.507783, -113.592490, 2019, on trunk of Ulmus
americana, M. Villeneuve & M. Lewis s.n. [UoA-CC-7] (hb. Haughland); Edmonton, Urban Monitoring
Site 107E, 53.548249, -113.617818, 2019, on trunk of Elaeagnus angustifolia, D. Haughland s.n. [UoA-
CC-9] (hb. Haughland); Edmonton, Urban Monitoring Site 149E, 53.587099, -113.610922, 2019, on trunk
79
of Salix cf. pentandra, D. Haughland & L. Hjartarson s.n. [UoA-CC-100] (hb. Haughland); Edmonton,
Urban Monitoring Site 37x, 53.494526, -113.597236, 2019, on trunk of Ulmus americana, M. Villeneuve &
M. Lewis s.n. [UoA-CC-115] (hb. Haughland); Woodcroft, Air Quality Monitoring Station, 53.563708, -
113.563508, 2019, on trunk of Prunus virginiana, D. Haughland & A. Hood s.n. [UoA-CC-107] (hb.
Haughland); Edmonton, Sir Wilfrid Laurier Park, 53.50834, -113.560926, 2019, on fallen Populus twigs,
D. Haughland 2019-121C & P. Williams (hb. Haughland).
GROUP 5. ORANGE LECANORINE CRUSTOSE LICHENS
Three species. Key literature: Arup 2009; Björk 2013; Brodo 2016; McCune 2017a, 2017b; Šoun
et al. 2011; Wetmore 2001, 2007a, 2007b. Additional species from this group may be present in Edmonton
but existing collections are too sparse to be confident. Identification of these three species is possible using
morphology and substrate.
1a. Growing on rock or concrete; all tissues orange, never grey .................................. Caloplaca feracissima
1b. Growing on bark or wood; thalline tissues may be grey or orange ........................................................... 2
2a. Thalline exciple orange, proper exciple often visible ................................................... Caloplaca pyracea
2b. Thallus exciple thick and grey, proper exciple indistinct/not visible .............................. Caloplaca cerina
Caloplaca pyracea (Ach.) Zwackh
(≡ Athallia pyracea (Ach.) Arup, Frödén & Søchting) FIGURE 25 C-D.
Common river valley, parkland, and tableland epiphyte. In Alberta records of this species were
historically included with Caloplaca holocarpa, but Arup (2009) considers C. holocarpa almost
exclusively saxicolous. Edmonton material: thallus pale grey to greyish orange-yellow, thin, inconspicuous,
or moderately-areolate. Apothecia abundant, scattered to crowded, usually flat to slightly convex, round to
irregular, 0.3–1.0 mm diameter, orange disk, thalline exciple slightly lighter orange than disk with proper
exciple visible and slightly raised or level with disk. Spores hyaline, uniseptate, polarilocular, ellipsoidal,
9–15 × 5–9 µm, septum 3-6 µm wide, 8 per ascus. Common across many deciduous tree species.
Chemistry: orange tissues K+ purple, C-, PD-, UV-, parietin, ± traces of fallacinal, emodin, teloschistin,
and parietinic acid (Arup 2009). Molecular data: no new sequences generated, genus-level and limited
species-level support in Arup et al. (2013).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 20, 53.464867, -113.505729, 2019, on trunk of Populus, D. Haughland & M. Cao s.n. [UoA-CC-33]
(hb. Haughland); Edmonton, Urban Monitoring Site 74, 53.548831, -113.592412, 2019, on trunk of
Populus, M. Lewis & M. Villeneuve s.n. [UoA-CC-99] (hb. Haughland); Edmonton, Urban Monitoring Site
83, 53.559556, -113.642705, 2019, on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n.
[UoA-CC-34] (hb. Haughland); Edmonton, Urban Monitoring Site 149E, 53.587099, -113.610922, 2019,
on trunk of Salix cf. pentandra, D. Haughland & L. Hjartarson s.n. [UoA-CC-100] (hb. Haughland);
Woodcroft, Air Quality Monitoring Station, 53.563693, -113.563604, 2019, on trunk of Prunus virginiana,
D. Haughland & A. Hood s.n. [UoA-CC-35] (hb. Haughland); Edmonton, Urban Monitoring Site 194E,
53.63778, -113.52964, 2019, on trunk of Fraxinus, D. Thauvette & M. Cao s.n. [UoA-CC-61] (hb.
Haughland); Edmonton, Sir Wilfrid Laurier Park, 53.50834, -113.560926, 2019, on bark of large downed
Populus, D. Haughland 2019-122B & P. Williams (hb. Haughland); Edmonton, 180 St and 99 Ave, 1976,
on twigs and dead branches, D.C. Lindsay s.n. (PMAE-B77.24.75); Edmonton, Terwillegar Park, 1977, on
rotting wood, D.C. Lindsay, s.n. (PMAE-B77.24.85).
Caloplaca cerina (Ehrh. ex Hedwig) Th. Fr. FIGURE 25 E-F.
Occasional river valley, parkland, and tableland epiphyte. This species can be confused with
Caloplaca pyracea, but C. cerina is rarer, has a thicker grey thalline margin, and lacks a visible orange
proper exciple. Edmonton material: thallus grey (pale to slate to dark grey), immersed to well-developed,
areolate or continuous. Apothecia single or clustered, 0.3–1.0 mm diameter, thalline exciple similar in color
to thallus (pale to dark grey), disk yellow to orange, sometimes white pruinose, proper exciple not visible.
Spores hyaline, uniseptate, polarilocular, 10–18 × 6–9 µm, septum 3–7 µm wide, 8 per ascus. Chemistry:
orange tissues K+ purple, C-, PD-, UV-, parietin, fallacinal, and teloschistin by TLC (Wetmore 2007a).
Molecular support: no new sequences generated, limited species level support (Frolov et al. 2021).
80
Figure 25. Orange lecanorine crustose Caloplaca s.l. of Edmonton. A-B, C. feracissima, Haughland 2020-
95. A, Apothecia on old concrete. B. Polarilocular spores with relatively narrow septa. C-D, C. pyracea. C,
Apothecia on Fraxinus pennsylvanica with proper exciple visible, surrounded by Phaeophyscia orbicularis,
UoA-CC-31. D, Polarilocular spores with wide septa, UoA-CC-45. E-F, C. cerina, UoA-CC-45. E,
Pruinose apothecia on Populus tremuloides, F, Apothecial cross-section showing thick thalline exciple.
81
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 40,
53.493796, -113.504715, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-
CC-8] (hb. Haughland); Edmonton, Urban Monitoring Site 62, 53.520757, -113.432138, 2019, on trunk of
Ulmus americana, S. Toni & A. Hood s.n. [UoA-CC-28] (hb. Haughland); Edmonton, Urban Monitoring
Site 62, 53.520926, -113.432139, 2019, trunk of Ulmus americana, D. Royko & D. Fielder s.n. [UoA-CC-
15] (hb. Haughland); Edmonton, Urban Monitoring Site 107E, 53.548554, -113.617129, 2019, on trunk of
Salix, D. Haughland s.n. [UoA-CC-97] (hb. Haughland); Edmonton, Urban Monitoring Site 23E,
53.470636, -113.620232, 2019, on trunk of Populus tremuloides, D. Haughland s.n. [UoA-CC-45] (hb.
Haughland); Edmonton, East Air Quality Monitoring Station, 53.548202, -113.367545, 2019, on trunk of
Populus, D. Thauvette & J. Birch s.n. [UoA-CC-6] (hb. Haughland); Edmonton, Sir Wilfrid Laurier Park,
53.50834, -113.560926, 2019, on bark of large downed Populus, D. Haughland 2019-122A & P. Williams
(hb. Haughland); Edmonton, Terwillegar Park, 1977, on rotting wood, D.C. Lindsay s.n. (PMAE-as minor
component of B77.24.85); 180 St. and 99 Ave., Edmonton, 1976, on twigs and dead branches, D.C.
Lindsay s.n. (PMAE-as minor component of B77.24.75).
Caloplaca feracissima H. Magn.
(≡ Xanthocarpia feracissima (H. Magn.) Frödén, Arup & Søchting) FIGURE 25 A-B.
Anthropogenic saxicole. Given the inconspicuous to absent primary thallus, the small apothecia,
spore size, and septum:length ratio, our material falls within the Caloplaca crenulatella complex. This
poorly resolved complex may represent a suite of phenotypically similar species (Arup 2009; McCune
2017b; Vondrák et al. 2011, 2017). Edmonton material: apothecia single to clustered, 0.2–0.5 mm diameter,
disk dull or dingy orange, proper and thalline exciple visible, orange. Spores simple, hyaline, 12–16 × 5–6
µm, septum 2.5–4 µm wide, 8 per ascus. Chemistry: orange tissues K+ purple, C-, PD-, UV-, secondary
metabolites not investigated. Molecular support: a single ITS sequence (isolate DLH33 from Haughland
2020-95A) is 99% percent identical to two Xanthocarpia feracissima GenBank sequences (MK110661, 4
positions different, 570 bp overlap, Kantor et al. 2018 unpublished; KC179129, U.S.A., 2 positions
different, 497 bp overlap, Arup et al. 2013). A sequence of uncultured fungus from house dust and indoor
air collected in Kansas City, U.S.A., was also 99% identical (KF800113, 4 positions different, 571 bp
overlap, Rittenour et al. 2014), possible evidence of propagules in other urban environments.
Specimen examined. – CANADA. ALBERTA: Edmonton, Spruce Avenue neighborhood,
53.563562, -113.498138, 2020, on old concrete sidewalk, D. Haughland 2020-95A (hb. Haughland).
GROUP 6. LECANORA S.L.
Eight species, five described here. See Group 2 for the biatorine Lecanora symmicta, Group 7 for
sorediate L. impudens and L. stanislai. Key literature: Björk 2013; Brodo 1984, 2016; McCune 2017a,
2017b; Śliwa 2007. Given recent taxonomic flux (Kondratyuk et al. 2019; Zhao et al. 2016) we retain
Lecanora in the broad sense and provide updated synonyms. Given the diversity of this genus, we
anticipate additional species will be found in Edmonton. Spot tests and examination of sections and spores
at high magnification and with polarized light is required for confident identification of most species.
1a. Apothecia biatorine or apothecia typically lacking and thallus composed of soredia ............................... 2
1b. Apothecia lecanorine, thallus immersed or episubstratic and areolate to subfoliose ................................. 3
2a. Apothecia biatorine ......................................................................................................... see Group 2 Key
2b. Apothecia typically lacking and thallus composed of soredia ......................................... see Group 7 Key
3a. Growing on concrete or rock; thallus well-developed, subfoliose ..........................Protoparmelia muralis
3b. Growing on bark or wood; thallus never subfoliose, at most areolate, often immersed ............................ 4
4a. Thalline exciple PD+ orange to red, best tested on apothecia cross-sections ............... Lecanora pulicaris
4b. Thalline tissues PD- .................................................................................................................................. 5
5a. Apothecial rim grey; spores 12-16 per ascus; primary thallus often visible as distinct white stain or
verrucae .............................................................................................................................. Lecanora sambuci
5b. Apothecial rim white; spores 8 per ascus; primary thallus typically lacking or indistinct ........................ 6
82
6a. Apothecia typically lacking pruina, polarizing crystals sparse in cross-sections ........................................
..................................................................................................................................... Lecanora cf. persimilis
6b. Apothecia typically pruinose, pruina, epihymenium and thalline exciple typically with abundant
polarizing crystals in sections .......................................................................................... Lecanora cf. hagenii
Lecanora cf. hagenii (Ach.) Ach.
(≡ Polyozosia hagenii (Ach.) S.Y. Kondr., Lőkös & Farkas, ≡ Myriolecis hagenii (Ach.) Śliwa, Zhao
Xin & Lumbsch) FIGURE 26 B.
Tableland, parkland, river valley and ravine system epiphyte. Common and variable on a diversity
of deciduous trees. Edmonton material: thallus immersed to indistinct, white to grey. apothecia pruinose
(immature and mature), at maturity with raised, white, crenulate rim. Apothecia 0.3–0.9 mm diameter,
crowded. Spores hyaline, simple, 10–12.5 × 5 µm, 8 per ascus. Epihymenium brown, hymenium hyaline,
hypothecium hyaline. Pruina, epihymenium and thalline exciple POL+. Epihymenium/hymenium C-.
Chemistry: all spot tests on thallus and thalline exciple negative, no secondary metabolites detected by TLC
(Ryan et al. 2004). Molecular support: unresolved. Sequences including the single sequence generated here
(isolate DLH15 from UoA-CC-61) fall into four distinct clades within the Lecanora dispersa group. Given
the uncertainty around the status of L. hagenii s.s. (see results section), we can confirm only that our
sequence is within the highly supported L. dispersa group, and that it is distinct from our single sequence of
L. cf. persimilis.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 31, 53.477955, -113.494143, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & A. Stordock
s.n. [UoA-CC-29] (hb. Haughland); Edmonton, Urban Monitoring Site 83, 53.559591, -113.642062, 2019,
on trunk of Ulmus americana, D. Haughland s.n. [UoA-CC-160] (hb. Haughland); Edmonton, Urban
Monitoring Site 111, 53.587075, -113.457605, 2019, on trunk of Fraxinus pennsylvanica, S. Toni & A.
Hood s.n. [UoA-CC-96] (hb. Haughland); Edmonton, Urban Monitoring Site 117, 53.600887, -113.501394,
2019, on trunk of Ulmus pumila, D. Haughland & M. Cao s.n. [UoA-CC-63] (hb. Haughland); Edmonton,
Urban Monitoring Site 194E, 53.63778, -113.52964, 2019, on trunk of Fraxinus, D. Thauvette & M. Cao
s.n. [UoA-CC-61] (hb. Haughland); Edmonton, Urban Monitoring Site 2E, 53.438762, -113.557303, 2019,
on trunk of Ulmus americana, L. Hjartarson & D. Haughland s.n. [UoA-CC-94] (hb. Haughland).
Lecanora cf. persimilis (Th. Fr.) Arnold
(≡ Polyozosia persimilis (Th. Fr.) S.Y. Kondr., Lőkös & Farkas, ≡ Myriolecis persimilis (Th. Fr.)
Śliwa, Zhao Xin & Lumbsch) FIGURE 26 D.
Occasional tableland epiphyte. Separated (with difficulty) from Lecanora cf. hagenii by scarcity of
pruina on even immature apothecia. Edmonton material: thallus immersed or lacking, apothecia sessile,
matte to slightly shiny, plane to slightly convex, with thin thalline margin, appearing almost biatorine,
diameter to 0.86 mm. Disk peach brown to medium brown, epihymenium brown in apothecia cross-
sections. Trebouxioud algae patchy, present as a thick layer in part below hypothecium. Spores 8–14 × 5.5–
6 µm, simple, 8 per ascus. Paraphyses wider at tips and sparsely branching. Very sparse polarizing crystals
present, scattered in hymenium and thalline exciple, lacking epipsamma. Chemistry: All spot tests negative,
no secondary metabolites detected by TLC (Ryan et al. 2004). Molecular data: no sequences in GenBank.
A single ITS sequence generated for this study (isolate DLH16 from UoA-CC-62) clustered in a clade
previously hypothesized to represent L. hagenii s.s. (Śliwa et al. 2012). Given the uncertainty around the
status of L. hagenii s.s. (see Results), we can confirm only that our sequence is within the highly supported
L. dispersa group, and that it is distinct from our single sequence of L. cf. hagenii.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 17,
53.466785, -113.5710074, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n.
[UoA-CC-62] (hb. Haughland); Edmonton, Urban Monitoring Site 62, 53.520926, -113.432139, 2019, on
trunk of Ulmus americana, D. Royko & D. Fielder s.n. [UoA-CC-15] (hb. Haughland); Edmonton, Urban
Monitoring Site 150E, 53.590394, -113.588685, 2019, on trunk of Ulmus americana, D. Haughland & L.
Hjartarson s.n. [UoA-CC-27] (hb. Haughland).
83
Figure 26. Thalli of Lecanora s.l. with 8-spored asci, Edmonton. A, L. pulicaris on fallen Betula
papyrifera, Haughland 2020-42. B, L. cf. hagenii on Ulmus pumila, UoA-CC-63. C, Protoparmeliopsis
muralis on top of concrete retaining wall, Haughland 2020-96. D, L. cf. persimilis on Ulmus americana,
UoA-CC-15.
Lecanora pulicaris (Pers.) Ach. FIGURE 26 A.
River valley epiphyte. Edmonton material: thallus white, appearing as a shiny stain on bark,
incipient apothecia in thalline verrucae. Apothecia with visible proper and white thalline exciple, disk red-
brown to pale brown, epruinose, to 1.2 mm diameter, constricted at base. Thalline exciple with cortex 13–
20 µm thick near edge, to >30 µm thick near base, with sparse, large, polarizing clumps of crystals, PD+
red, K+ yellow to orange, C-, UV- in section. Epihymenium tan with abundant tiny polarizing granules
throughout, PD-, C-, K+ yellow in section. Hymenium and subhymenium hyaline. Spores globose to
ellipsoidal, 8–13 × 5–10 µm, hyaline, non-septate, 8 per ascus. Chemistry: thallus PD+ orange or PD-, K+
yellow, KC-, C-, UV-, atranorin, ± chloroatranorin (trace), confumarprotocetraric acid (trace),
fumarprotocetraric acid (major), roccellic acid (major) (Ryan et al. 2004). Molecular support: no new
sequences generated, limited phylogenetic support for this species in Malíček et al. (2017) and Lee & Hur
(2020). Specimens examined. – CANADA. ALBERTA: Edmonton, Patricia Ravine, 53.503105, -
113.592863, 2020, on bark of recently downed Betula papyrifera, D. Haughland 2020-42 & A. Hood (hb.
Haughland); Edmonton, University of Alberta North Campus, 53.5253952, -113.525717, 2021, on Pinus
mugo, J. Singh & K. Schafer s.n. [UoA-CC21-18] (hb. Haughland)
84
Figure 27. Lecanora sambuci morphology and anatomy, Edmonton, UoA-CC-101. A, Thallus on Fraxinus
pennsylvanica. B, Asci with 12-16 simple spores. C, Apothecia cross-sections under white light. D,
Apothecia cross-sections under polarized light showing crystals in thalline exciple.
*Lecanora sambuci (Pers.) Nyl.
(≡ Polyozosia sambuci (Pers.) S.Y. Kondr., Lőkös & Farkas, ≡ Myriolecis sambuci (Pers.) Clem.)
FIGURE 27.
Tableland and parkland epiphyte. The type species of the recently resurrected genus Myriolecis
(Zhao et al. 2016), formerly referred to as the L. dispersa group, these are the first records for Alberta. The
species has been recorded from both neighboring provinces as well as states to the south (CNALH 2020,
Thomson 1997). Edmonton material: apothecia clustered, brown epruinose disk with prominent grey
smooth to coarse thalline rim. Spores hyaline, simple, 7–8.5 × 3–5 µm, 12–16 per ascus, asci Lecanora-
type with K/I+blue thollus and non-staining central axis. Thalline exciple with polarizing crystals.
Chemistry: all spot tests negative, no secondary metabolites detected by TLC. Molecular support: none, one
ITS sequence in GenBank, no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 11,
53.450690, -113.474695, 2019, on trunk of Fraxinus pennsylvanica, D. Thauvette & J. Wasyliw s.n. [UoA-
CC-101] (hb. Haughland); Edmonton, Urban Monitoring Site 36, 53.494916, -113.620094, 2019, on trunk
of Fraxinus pennsylvanica, D. Haughland & L. Hjartarson s.n. [UoA-CC-113] (hb. Haughland);
Edmonton, Urban Monitoring Site 185E, 53.62906, -113.56883, 2019, on trunk of Populus, D. Thauvette &
M. Cao s.n. [UoA-CC-91] (hb. Haughland).
85
Protoparmeliopsis muralis (Schreber) M. Choisy
(≡ Lecanora muralis (Schreber) Rabenh.) FIGURE 26 C.
Anthropogenic saxicole. A distinctive placodioid species, we recorded a single specimen from
concrete. Edmonton material: appressed flat to slightly concave-lobed thallus with pale greenish-yellow
upper cortex and blue-black lobe edges. In the transverse section of the lobes, hyphae of the upper cortex
can be seen regularly invading down through the algal layer. Similarly, hyphae of the thalline exciple can
be seen invading the algal layer below the hypothecium in apothecia cross-sections. The apothecia were
clustered centrally, characterized by their tan to peach-brown epruinose disk and broad, flat thalline
exciple. Epihymenium tan with fine polarizing crystals that largely dissolved in K, hyaline hymenium to 70
µm thick. Paraphyses similar in width through their length, regularly septate, branching near the base. Asci
8-spored, spores hyaline, simple, sub-globose to ovoid, 9–11 × 5–6 µm. Thalline exciple with polarizing
crystals. Chemistry: all spot tests are negative in the medulla; the upper cortex is K-, C-, KC+ yellow in
older regions, PD-, UV365-, UV254-. Secondary metabolites detected by TLC: usnic acid, zeorin, leucotylin,
unknown fatty acids. Molecular support: strong genus and species-level support (Zhao et al. 2016, Tripp et
al. 2019), no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Spruce Avenue Neighborhood,
53.562815, -113.497646, 2020, on top of old, low concrete retaining wall, D. Haughland 2020-96 & K.
Tichkowski (hb. Haughland).
GROUP 7. SOREDIATE/GRANULAR CRUSTOSE LICHENS
Five species. Key literature: Brodo 1984, 1991; Lendemer 2010, 2011, 2013; Lendemer &
Hodkinson 2013; Malíček 2014; Malíček et al. 2017; McCune 2017a, 2017b; Vondrák et al. 2009. This
group is undoubtedly underrepresented in our study, and we cannot say whether most species are rare or
common with any certainty. Ochrolechia arborea is an additional species we could not confirm. Brodo
(1991) reported this species from nearby Wagner Natural Area, which we’ve excluded geographically from
this study; Elsinger et al. (2007) also reported it from the industrial zone east of Edmonton. The
identification of some species remains uncertain, and TLC is necessary to distinguish others.
1a. Thallus initially episubstratic and smooth to areolate, forming discrete soralia ........................................ 2
1b. Thallus formed entirely of soredia, no cortex present ............................................................................... 3
2a. Orange areolate thallus on anthropogenic substrates, with marginal soralia ................. Caloplaca tominii
2b. Greyish episubstratic thin to verrucose thallus on trees, forming round, laminal soralia ............................
.......................................................................................................................................... Lecanora impudens
3a. Placodioid, soredia embedded in a deep (to 1 cm thick) hypothallus, resembling drier lint, PD+ orange ..
...................................................................................................................................................Lepraria finkii
3b. Discrete soredia not embedded in hypothallus, at most 1 mm thick, PD- ................................................. 4
4a. Atranorin and 2 orange fluorescing unknowns in 365nm light (TLC) ..............................Lecidella albida
4b. Usnic acid, zeorin (TLC) .............................................................................................. Lecanora stanislai
Caloplaca tominii (Savicz) Ahlner
(≡ Xanthocarpia tominii (Savicz) Frödén, Arup & Søchting) FIGURE 28 F.
Anthropogenic terricole/saxicole. In the most inclusive treatment of sorediate Caloplaca s.l.
(Vondrák et al. 2009), our material keyed to the C. citrina (Hoffm.) Th. Fr. group (≡ Flavoplaca citrina
(Hoffm.) Arup, Frödén & Søchting), specifically C. austrocitrina Vondrák, Říha, Arup & Søchting (Arup
2006). However, ITS showed our material to belongs to C. tominii, reinforcing the difficulty in accurately
identifying infertile Caloplaca s.l., particularly within currently accepted genera. The only other Alberta
record of this species is from the Rocky Mountains Natural Region (Wetmore 2001). Edmonton material:
thalli areolate, attached firmly throughout, to 0.5–2.0 mm in diameter and 180–290 µm thick in cross-
section. Areoles are yellow, with soralia developing marginally. In larger thalli, soralia appear to first
develop on the margins of the upper surface and then towards the center, occasionally becoming covered
86
Figure 28. Sorediate crustose lichens of Edmonton, plate 1 of 2. A-B, Lecidella albida, Haughland 2020-
28. A, Thallus on Picea bark flake. B, Granules in water mount. C-D, Lepraria finkii, Haughland 2020-21.
C, Vertical section showing thick hypothallus. D, Granules with protruding hyphae and persistent crystals
after treatment with K, in polarized light. E, Lecanora impudens, UoA-CC-51. F, Caloplaca tominii,
Haughland 2020-97A.
87
in granular soredia, apothecia occasional, but no mature asci or spores found. Apothecial thalline rim
sorediate. See Wetmore (2001) for detailed comparison with other sorediate Caloplaca. Chemistry:
medulla and cortex K+ purple, parietin, fallacinal, emodin and teloschistin by TLC (Wetmore 2007a).
Molecular support: a single ITS sequence (isolate DLH7 from Haughland 2020-97A) was most similar in
megablast to C. tominii specimens from Europe and Asia (Genbank accessions HQ69950, MG954185,
HQ699626, query cover 87–97%, percent identity 96–99%). No accessioned C. citrina or C. austrocitrina
exceeded 87% percent identity despite similar query cover (92–97%).
Specimens examined. – CANADA. ALBERTA: Edmonton, Spruce Avenue neighborhood,
53.563562, -113.498138, 2020, on old concrete sidewalk, D. Haughland 2020-95B (hb. Haughland), on
weathered and mossy indoor-outdoor carpet, D. Haughland 2020-97A (hb. Haughland).
Lecanora impudens Degel. FIGURE 28 E.
Predominantly river valley and parkland epiphyte. A relatively common species described as
having a thin, grey episubstratic thallus with irregular to more or less circular excavate soralia that produce
granular soredia (Brodo 1984, Lendemer 2010). We used two specimens identified by I. Brodo in PMAE as
our morphological benchmark; they were similarly suggestive of Pertusaria in morphology and color
(Moose Factory Island, Ontario, 1969, bark of Populus balsamifera, 51.25, -80.63, Brodo 14672; Boulder,
Colorado, 1964, bark of fallen spruce, Fourth of July Campground 8 miles NW of Nederland, Sierk 2479).
Edmonton material: thalli typically continuous, episubstratic, grey to grey-yellow, most forming verrucae
that dissolved apically into soralia. Soralia varied from discrete, circular, and excavate with granular to
powdery white soralia (as in the specimens cited above) to irregular, crowded, but not becoming
completely confluent due to persistence of soralia thalline rims. In the latter specimens the granular soredia
were greenish-white in fresh material, cream to pale yellow in older collections. The soralia became
excavate as they enlarged and towards the center of the thallus. The specimen on Sorbus was unique; it had
sparse, poorly formed, almost globular apothecia. No mature asci or spores were found, and the thalline
exciple was allophana-type (following Brodo 1984). Photobiont trebouxioid. Chemistry: PD-, C-, K+
yellow, KC-, UV365+ dull white, UV254+ dull orange. Secondary metabolites by TLC: atranorin, ±
unknowns including trace fatty acid (Rf 3/4/?), unknown blue-fluorescing trace compound (Rf 4/5–6/6).
Atranorin is the only metabolite common across the literature, with occasional additional fatty acids or
terpenoids noted in some reports (Brodo 1984, Malíček 2014). Molecular support: this species is difficult to
discriminate from Lecanora allophana f. sorediata Vain., and the limited molecular evidence to date
suggests they may be conspecific (Malíček et al. 2017). More work is needed.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, McKinnon Ravine,
1976, on tree bark, D.C. Lindsay s.n. (PMAE-B77.24.152), D.C. Lindsay s.n. (PMAE-B77.24.170);
Edmonton, Urban Monitoring Site 185E, 53.62988, -113.56831, 2019, trunk of Fraxinus, D. Thauvette &
M. Cao s.n. [UoA-CC-51] (hb. Haughland); Edmonton, Urban Monitoring Site 28, 53.479694, -
113.549752, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n. [UoA-CC-52] (hb.
Haughland); Edmonton, Urban Monitoring Site 47, 53.505933, -113.552966, 2019, on trunk of Populus
balsamifera, D. Haughland s.n. [UoA-CC-47] (hb. Haughland); Edmonton, Urban Monitoring Site 68,
53.530342, -113.553482, 2019, on trunk of Ulmus, D. Haughland s.n. [UoA-CC-11] (hb. Haughland);
Edmonton, Urban Monitoring Site 86, 53.561366, -113.548002, 2019, on trunk of Ulmus americana, D.
Haughland & A. Hood s.n. [UoA-CC-50] (hb. Haughland); Edmonton, Urban Monitoring Site 148E,
53.587522, -113.640605, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n.
[UoA-CC-48,125] (hb. Haughland); Edmonton, Urban Monitoring Site 23E, 53.470689, -113.620142,
2019, on trunk of Populus tremuloides, D. Haughland s.n. [UoA-CC-49] (hb. Haughland).
*Lecanora stanislai Guzow-Krzemińska, Lubek, Malíček & Kukwa FIGURE 29.
River valley epiphyte. Leprarioid, sterile lichens containing usnic acid and zeorin are poorly
understood in North America (Lendemer & Hodkinson 2013). Species in this group include Lecanora
expallens Ach., a variable, typically sterile lichen that contains thiophanic acid as a major metabolite (Ryan
et al. 2004); the latter may be in low concentration in shaded microhabitats and difficult to detect with TLC
(Guzow-Krzemińska et al. 2017). Lecanora thysanophora R.C. Harris, common in eastern North America
and present in the pacific northwest (J. Lendemer, pers. comm.), typically has a well-developed fibrous
prothallus and additional compounds such as porphyrillic acid and unknown substances (Harris et al. 2001).
88
Figure 29. Sorediate crustose lichens of Edmonton, plate 2 of 2. Lecanora stanislai, growing on Picea
glauca snags, Haughland 2020-103. A, Thallus. B, Close-up of aggregate soredia/granules. C, Soredia in
water mount showing trebouxioid algae and knobby, septate hyphae. D, Soredia under polarized light,
showing fine polarizing crystals prior to treatment with K.
Lecanora compallens Herk & Aptroot, described from Europe and not known from North America, has
soralia that start as punctiform openings in a thin to verrucose thallus and medulla is evident below the
soredia (van Herk and Aptroot 1999, Guzow-Krzemińska et al. 2017). The saxicolous Lithocalla ecorticata
(J.R. Laundon) Orange known from Europe (Orange 2021, Smith et al. 2009) was excluded from the North
American flora by Lendemer & Hodkinson (2013), and reports of zeorin are now attributed to
contamination (Orange et al. 2017, Orange 2021). Lendemer & Hodkinson (2013) described Leprocaulon
knudsenii Lendemer & Hodkinson, a saxicolous leprarioid species with usnic acid and zeorin from southern
California, but they lacked adequate material to describe additional, phenotypically similar but molecularly
distinct collections that belong to Lecanora, as did Guzow-Krzemińska et al. (2017). More recently,
Lecanora stanislai was described from Europe, Asia, and western North America including British
Columbia (Guzow-Krzemińska et al. 2017). The morphological description is the best fit for our material.
Edmonton material: extensive, thin granular aggregate varying from pale blue-green to cream in color,
granules 40–50 µm in diameter, over a thin, white episubstratic prothallus. Hypothallus and delimited
soralia lacking and granules largely without projecting hyphae except in senescing, cream-colored patches.
Photobiont trebouxioid, cells 9–12 µm in diameter, each algal cell surrounded by a single layer of knobby,
septate, branching hyphae, each hyphal segment 2–3 × 3–7 µm. Granules with fine POL+ crystals that
dissolve entirely in K. Cream-colored granules contain dead, cavitated algal cells. No apothecia or pycnidia
found except black apothecia of an unidentified, apparently non-lichenized ascomycete fungus. Chemistry:
granules PD-, K+ pale yellow, KC-, C-, UV365-, UV254-. Secondary metabolites by TLC: usnic acid, zeorin
89
(trace). Molecular support: currently lacking for Edmonton collections, two samples failed to amplify
cleanly enough to sequence. See Guzow-Krzemińska et al. (2017) for phylogenetic placement.
Specimen examined. – CANADA. ALBERTA: Edmonton, along Rat Creek, Kinnaird Ravine,
53.558953, -113.459253, 2020, on trunks of Picea glauca snags, D. Haughland 2020-103 (hb. Haughland).
**Lecidella albida Hafellner FIGURE 28 A-B.
Ravine system epiphyte. Molecular data was necessary to place this collection, as there are sterile
crusts that produce atranorin in multiple genera including Bacidia, Cliostomum, Lecanora, and Lecidella.
Edmonton material: on the flaky bark of the trunk of a slightly leaning Picea snag, on the snow-collecting
upward surface, growing with Candelaria pacifica and Ramalina. Thallus consists of a thin layer of pale
greenish-yellow discrete granules (aggregate-type, sensu Lendemer 2011) that start as larger granules and
dissolve into a thin layer of smaller granules over a pale grey prothallus. Granules are discrete bundles of
hyphae around green trebouxioid algae measuring 7–9 µm in diameter that lack projecting hyphae or a
Lepraria-like hypothallus. Abundant POL+ fine crystals that dissolve with K coat the bundles. Chemistry:
K+ pale yellow, PD-, C-, KC-, UV365+ dull orange, UV254-. Secondary metabolites detected by TLC:
atranorin, two possible xanthones that fluoresce orange under UV365. Molecular support: A single ITS
sequence (isolate DLH1 from Haughland 2020-28) forms a well-supported clade basal to the clades of
Lecidella sequenced to date, with a sequence of L. albida from Switzerland (GenBank Accession
KX132964; Fig. 6 herein). Additional work is required to be confident these specimens belong in the genus
Lecidella.
Specimen examined. – CANADA. ALBERTA: Edmonton, along Rat Creek, Kinnaird Ravine,
53.558953, -113.459253, 2020, on trunk of Picea glauca snag, D. Haughland 2020-28 (hb. Haughland).
Lepraria finkii (B. de Lesd.) R.C. Harris FIGURE 28 C-D.
River valley and ravine system epiphyte. Edmonton material: placodioid (sensu Lendemer 2011),
consisting of granules embedded in a thick hypothallus. Up to 1 cm thick, the pale green-blue granules have
abundant, septate, hydrophobic, projecting hyphae 3–4 µm in diameter that branch regularly, with globose
green algae 6 µm in diameter. Abundant POL+ crystals that persist after treatment with K coat the outside
of the hyphae. Chemistry: K+ pale yellow, PD(ethanol)+ slow orange, UV365+ dull white, UV254-.
Secondary metabolites detected by TLC: atranorin, zeorin, stictic acid (major or in trace amounts), ±
norstictic acid (trace). Molecular support: weak. Using megablast, our single sequence of Lepraria finkii
(isolate DLH9 from Haughland 2020-100) scored highest with accessioned L. finkii sequences; however,
despite high query coverage (>90%), the percent identities were ≤75%. For example, our ITS sequence
differed by 130 positions from GenBank Accession MK629287 (Bolivia, 504 bp overlapping range, 75%
percent identity, Barcenas-Peña et al. 2021). Because our sole sequence contained a relatively high number
of ambiguous positions, and there are relatively few ITS sequences for comparison, we refrained from
further phylogenetic analyses.
Specimens examined. – CANADA. ALBERTA: Edmonton, Hawrelak off-leash trail, 53.520696,
-113.541533, 2020, on tree base, D. Haughland 2020-21 (hb. Haughland); Edmonton, along Rat Creek,
Kinnaird Ravine, 53.558953, -113.459253, 2020, on trunk of Picea glauca snag, D. Haughland 2020-100
(hb. Haughland); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315, 2021, on decayed stump, D.
Haughland 2021-23B (hb. Haughland).
GROUP 8. ORANGE AND YELLOW FOLIOSE LICHENS
Thirteen species. Key literature: Brodo 2016; Brodo et al. 2001; Lindblom 1997; Stapper 2012.
1a. Lobes orange, K+purple cortex (anthraquinones) ..................................................................................... 2
1b. Lobes greenish yellow, pale yellow or bright yellow, K- .......................................................................... 8
2a. Rhizines present, white, often visible beyond lobe tips .............................................. 3 (Xanthomendoza)
2b. Rhizines lacking, may be attached with sparse, stubby, peg-like hapters hidden beneath lobes .................
................................................................................ 5 (Xanthoria s.l. including Polycaulionia & Rusavskia)
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3a. Esorediate; apothecia typically present; laminal pycnidia visible as orange bumps ....................................
................................................................................................................................ Xanthomendoza hasseana
3b. Sorediate; apothecia occasionally present; pycnidia mostly not apparent ................................................. 4
4a. Lobes appressed, relatively robust, with abundant marginal, crescent- or “bird nest”-shaped soralia
formed between the upper and lower cortex; ubiquitous ............................................. Xanthomendoza fallax
4b. Lobes ascending, thin, delicate, with labriform, terminal soralia; rare.................... Xanthomendoza fulva
5a. Thallus of narrow, subfruticose, ascending lobes; forming terminal blastidia/soredia, apothecia rare .......
................................................................................................................................... Polycauliona candelaria
5b. Thallus of flat, appressed lobes or cushion-like, ill-formed lobes; esorediate, apothecia common........... 6
6a. Thallus cushion-like with lumpy, ill-differentiated, lobes; on wood or bark ........ Polycauliona polycarpa
6b. Thallus of flat, appressed, clearly foliose lobes; on wood, bark or rock ................................................... 7
7a. Lobes broad, petal-like, concave; thallus appressed but apothecia in center of thallus becoming raised;
only records in Alberta are from ornamental trees and shrubs ......................................... Xanthoria parietina
7b. Lobes, narrow, tightly adnate to almost crustose; apothecia adnate to thallus; commonly found on rock
or on anthropogenic substrates ............................................................................................ Rusavskia elegans
8a. Lobes bright yellow ................................................................................................................................... 9
8b. Lobes pale to greenish yellow ................................................................................................................. 11
9a. Lobes relatively broad and large, ≥0.5 cm wide, ascending, with abundant, continuous marginal soralia
containing abundant powdery soredia ................................................................................. Vulpicida pinastri
9b. Lobes narrow and tiny, ≤0.3 mm wide; soralia restricted to lobe tips or rarely, most of the thallus
dissolving into coarse soredia .................................................................................................. 10 (Candelaria)
10a. Lobes appressed; lower surface corticate, white with well-developed white rhizines; relatively rare in
Alberta, more common in tableland habitats ............................................................. Candelaria concolor s.l.
10b. Lobes typically ascending; lower surface largely ecorticate and greenish, lacking well-developed
rhizines; common in Alberta, particularly in riparian and wetland habitats ..................... Candelaria pacifica
11a. Lobes small, narrow (width typically ≤2 mm), appressed and linear ................... Parmeliopsis ambigua
11b. Lobes large (typically wider than 1 cm), rounded, ruffled ......................................... 12 (Shield lichens)
12a. Upper cortex more grey than yellow; lower cortex near lobe tips beige to tan, with abundant short,
simple rhizines to the lobe edge; cortex K+ yellow, medulla PD- ...................................... Punctelia caseana
12b. Upper cortex more yellow than grey; lower cortex brown to black, rhizines sparse; cortex K-, KC+
yellow, medulla PD- or PD+ orange-red ................................................................ 13 (Yellow shield lichens)
13a. Pseudocyphellae absent; soralia laminal; medulla PD+ orange-red ..........................................................
............ Flavoparmelia caperata [If present, rare in Alberta; see discussion under Flavopunctelia flaventior]
13b. Pseudocyphellae present, sparse to abundant; soralia laminal and/or marginal; medulla PD-
.......................................................................................................................................... 14 (Flavopunctelia)
14a. Soralia marginal and laminal; pseudocyphellae obvious and up to 1 mm across, elongate and
branched, developing into laminal soredia .............................................................. Flavopunctelia flaventior
14b. Soralia primarily marginal, horseshoe-shaped, on suberect lobes; pseudocyphellae rare or inapparent
and punctiform; upper surface often with faint white angular maculae visible with magnification; .................
................................................................................................................................... Flavopunctelia soredica
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Figure 30. Yellow sorediate foliose lichens of Edmonton, plate 1 of 2: anatomy and morphology of the
minute Candelaria. A, C, E, C. concolor s.l. on Fraxinus pennsylvanica, UoA-CC-123. A, Colony showing
shade-grown thalli in part. C, Close-up of lobes with rhizines projecting beyond lobe tips. E, Transverse
lobe sections showing well-developed lower cortex and rhizines, under white (top) and polarized light
(bottom). B, D, F, C. pacifica on Picea twigs, Haughland 2021-32. B, Thallus. D, Close-up of lobes
showing ascending habit and blastidia-like soredia. F, Transverse lobe sections showing absence of lower
cortex and penetration of upper cortex through the medulla, under white (top) and polarized light (bottom).
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Candelaria concolor (Dickson) Stein s.l. FIGURE 30 A, C, E.
Tableland epiphyte. With the separation of Candelaria pacifica from C. concolor (Westberg &
Arup 2011), all historical Alberta collections require re-determination. Based on collections received to
date, C. concolor is rare in Alberta (ABMI 2020, unpub. data) so we were surprised to make multiple
collections within Edmonton. This is a tiny yellow foliose lichen with lobes that typically do not exceed 0.5
mm in width. It lies appressed to the substrate and forms terminal soralia; apothecia are rare. This species
differs from the more common C. pacifica in having a relatively well-developed white lower cortex and
white rhizines that are often visible around the edges of the lobes. If material is fertile, it has >30 spores per
ascus (vs. eight in C. pacifica). Edmonton material: lobes typically appressed, soralia more granular-
sorediate (vs. blastidiate in C. pacifica); see Stapper (2012) for excellent illustrations of both species. All
detections to date on boulevard trees (vs. C. pacifica, which is common within river valley parks).
Chemistry: K-, KC-, PD-, UV-, calycin and pulvinic dilactone by TLC (Westberg & Nash 2002).
Molecular support: a single ITS sequence (isolate DLH37 from UoA-CC-96) differed by four positions
from C. concolor GenBank Accession KT695365 (Ontario, Canada: 528 bp overlapping range, 99%
percent identity) and MK966426 (New York, USA: 497 bp overlap, 99% percent identity). Our phylogeny
places our material, along with eastern North American collections, in a clade distinct from C. concolor in
Europe, and sister to Candelaria asiatica from South Korea (Liu & Hur 2018) and China (Kondratyuk et al.
2020). Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 39,
53.494127, -113.525569, 2019, on trunk of Fraxinus pennsylvanica, D. Evans & S. Toni s.n. [UoA-CC-
116] (hb. Haughland); Edmonton, Urban Monitoring Site 60, 53.520974, -113.480625, 2019, on trunk of
Fraxinus pennsylvanica, D. Thauvette & J. Wasyliw s.n. [UoA-CC-82] (hb. Haughland); Edmonton, Urban
Monitoring Site 111, 53.587075, -113.457605, 2019, on trunk of Fraxinus pennsylvanica, S. Toni & A.
Hood s.n. [UoA-CC-96] (hb. Haughland); Edmonton, Urban Monitoring Site 135, 53.642026, -113.501278,
2019, on trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-CC-123] (hb. Haughland).
*Candelaria pacifica M. Westb. & Arup FIGURE 30 B, D, F.
Predominantly river valley and ravine system and parkland epiphyte. This is the first published
report of this species in Alberta, aside from publicly available data from the Alberta Biodiversity
Monitoring Institute (ABMI 2020). Common west of the Rocky Mountains, this tiny, variable species
forms bright yellow rosettes up to 5 mm wide, sometimes dissolving into blastidia with few visible lobes.
Individual lobes rarely exceed 0.4 mm wide, and they typically ascend off the substrate. The lower cortex is
poorly developed or lacking, the medulla thin. See Candelaria concolor for distinguishing traits.
Chemistry: K-, KC-, PD-, UV-, pulvinic acid, pulvic acid lactone, calycin detected by high performance
TLC, and vulpinic acid by high performance liquid chromatography (Westberg & Arup 2011). Molecular
support: no new sequences generated, the separation from C. concolor is well-supported in phylogenetic
analyses (Westberg & Arup 2011, Fig. 2 herein).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 3, 53.440376, -113.484487, 2019, on trunk of Ulmus americana, J. Birch & J. Wasyliw s.n. [UoA-CC-
200] (hb. Haughland); Edmonton, Urban Monitoring Site Beverly Air Quality Monitoring Station,
53.566860, -113.397464, 2019, on trunk of Ulmus americana, D. Thauvette & J. Birch s.n. [UoA-CC-138]
(hb. Haughland); Saskatchewan Drive off-leash area, in ravine, 53.52, -113.54, 2017, on lower dead twigs
of live Picea glauca, D. Haughland 2017-1 & P. Williams (hb. Haughland); Edmonton, MacKenzie
Ravine, 1976, on bark, D.C. Lindsay s.n. (PMAE-B77.24.159), D.C. Lindsay s.n. (PMAE-B77.24.168);
Edmonton, grassy park next to Saskatchewan Drive bordering river valley, 53.513197, -113.53866, 2021,
on Picea twigs, D. Haughland 2021-32 (hb. Haughland); Edmonton, MacKenzie Ravine, by boardwalk,
53.52914, -113.5603, 2020, on downed Picea glauca twigs at the edge of mineral seep, D. Haughland
2020-106B (hb. Haughland); Edmonton, River Loop Trail south of Fort Edmonton, 53.500627, -
113.576611, 2021, on Betula papyrifera, D. Haughland 2021-18 & S. Toni (hb. Haughland); Edmonton,
Mill Creek ravine, 53.517222, -113.473889, 2020, on Picea glauca, D. Haughland 2020-115A & P.
Williams (hb. Haughland).
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Figure 31. Yellow (to green-yellow) sorediate foliose lichens of Edmonton, plate 2 of 2: species with thalli
≥1 cm in diameter. A, Flavopunctelia flaventior on Betula, Haughland s.n. B, Flavopunctelia soredica on
Picea glauca twigs, Haughland 2020-106A. C, Vulpicida pinastri on Betula, Haughland unvouchered
observation. D, Parmeliopsis ambigua on lignin, Haughland 2021-2A.
Flavopunctelia flaventior (Stirton) Hale FIGURE 31 A.
Common river valley epiphyte, rare in tableland and parklands. Thalli can exceed palm-size, with
large, rounded, ruffled lobes with abundant, laminal pseudocyphellae, and typically granular soredia mostly
in laminal soralia (with a few in crescent-shaped marginal soralia). Some thalli are difficult to distinguish
from F. soredica which has less conspicuous, sparse pseudocyphellae and largely marginal soralia with
finer soredia. This genus has a dark-brown to black lower cortex with sparse, simple rhizines, which helps
distinguish it from similarly-sized Punctelia (grey-green upper cortex and pale, tan, or pale brown lower
cortex). Locally this genus often is confused with Flavoparmelia caperata, which is very rare or perhaps
absent in Alberta; it can be distinguished by the lack of pseudocyphellae on the upper surface and the PD+
red medulla. Chemistry: upper cortex KC+ yellow, medulla C+ red, KC+ red, PD-, all other tests negative,
Secondary metabolites detected by TLC: usnic acid and lecanoric acid. Molecular support: no new
sequences generated, the separation from F. soredica is well-supported in our phylogenetic analyses.
94
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Whitemud Park, near Alfred H. Savage Centre, 53.501791, -113.559168, 2020, on Betula, J. Bell
(unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/11785/details); Edmonton,
Kinsmen Park, 53.528793, -113.518148, 2019, epiphytic, T.L. Dueck (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/8155/details); Edmonton, MacKenzie Ravine, 1976, on
Populus wood, D.C. Lindsay s.n. (PMAE-B77.24.34); Edmonton, Stony Plain Rd. and 100 Ave. at 148 St.,
1976, on wooden footbridge, D.C. Lindsay s.n. (PMAE-B77.24.61); Edmonton, Emily Murphy Park, 1976,
on Betula bark, D.C. Lindsay, s.n. (PMAE-B77.24.91); Edmonton, Whitemud Park, 1976, on wood, D.C.
Lindsay s.n. (PMAE-B77.24.112 ); Edmonton, Terwillegar Footbridge, 53.483201, -113.600184, 2021, on
Betula papyrifera, D. Haughland 2021-11 (hb. Haughland); Edmonton, Hawrelak Trail off-leash park,
53.522361, -113.54378, 2020, on decorticate Picea twigs, D. Haughland 2020-25 (hb. Haughland).
Flavopunctelia soredica (Nyl.) Hale FIGURE 31 B.
River valley epiphyte. Similar to Flavopunctelia flaventior except F. soredica has primarily
marginal soralia giving rise to crescent-shaped, powdery lobe tips and less abundant pseudocyphellae. It is
also rarer than F. flaventior; see that species for comparisons with other shield lichens. Chemistry: upper
cortex KC+ yellow, medulla C+ red, KC+ red, all other tests negative. Secondary metabolites detected by
TLC: usnic acid and lecanoric acid. Molecular support: a single ITS sequence of Flavopunctelia soredica
(isolate DLH13 from Haughland 2020-16) is basal within a well-supported clade of F. soredica sequences.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Hawrelak Trail off-leash park, 53.520696, -113.541533, 2020, on Betula bark, D. Haughland 2020-16 (hb.
Haughland); Edmonton, Patricia Ravine, 53.502112, -113.592209, 2020, epiphytic, S. Toni (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/12492/details); Edmonton, MacKenzie
Ravine, 53.52914, -113.5603, 2020, on downed Picea glauca twigs at edge of mineral seep, D. Haughland
2020-106A (hb. Haughland; NatureLynx record https://naturelynx.ca/sightings/14351/details); Edmonton,
Hawrelak Park, 53.51665, -113.538811, 2020, on Betula papyrifera, D. Haughland 2020-113 (hb.
Haughland).
Parmeliopsis ambigua (Wulfen) Nyl. FIGURE 31 D.
River valley and ravine system lignicole. A common boreal species characterized by pale yellow,
appressed, narrow lobes (to 2 mm wide) with round laminal soralia. It can be confused with Parmeliopsis
hyperopta (Ach.) Arnold, a relatively common co-occurring species that differs in its whitish-grey
coloration (lacks usnic acid in the cortex); to date P. hyperopta has not been detected in Edmonton.
Chemistry: medulla UV+ blue-white, cortex KC+ yellow, all other tests negative, usnic acid, divaricatic
acid, nordivaricatic acid detected by TLC. Molecular support: monophyly confirmed by Tehler & Källersjö
(2001), no new sequences generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
53.502711, -113.60241, 2020, lignicolous on downed log, D. Haughland & C. Shier (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/14354/details); Edmonton, MacKenzie
Ravine, 1976, on wood, D.C. Lindsay s.n (PMAE-B77.24.35); Edmonton, Terwillegar Footbridge, 53.4797,
-113.594315, 2021, on Picea snag, D. Haughland 2021-2A (hb. Haughland).
Polycauliona candelaria (L.) Frödén, Arup, & Søchting
(≡ Xanthoria candelaria (L.) Th. Fr.) FIGURE 32 A.
Occasional river valley epiphyte. Thalli form small cushions typically less than the size of a
quarter, the bright-orange lobes are narrow, flattened, branched, and usually ascending to erect. Granular
soredia form at the lobe tips and along the lobe margins. Lacking rhizines, instead with small peg-like
haptors attaching proximal lobes to the substrate. Sometimes mistaken for Candelaria, this species can be
distinguished by the orange cortex and the K test. Chemistry: orange-pigmented parts K+ purple (vs. K- in
Candelaria), all other spot tests negative, parietin, fallacinal, emodin, teloschistin, and parietinic acid by
TLC (Lindblom 2004b). Molecular support: a variable species in need of phylogenetic work, no new
sequences generated.
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Figure 32. Orange foliose lichens of Edmonton, plate 1 of 2: native species. A, Polycauliona candelaria on
Picea twigs, Haughland 2021-34. B, Polycauliona polycarpa on Picea twigs, Haughland 2021-28. C,
Rusavskia elegans on metal bridge girders, Dueck https://naturelynx.ca/sightings/9806/details. D,
Xanthomendoza fallax on Populus, Haughland unvouchered observation. E, X. fulva on Ulmus americana,
UoA-CC-143. F, X. hasseana, Hjartarson https://naturelynx.ca/sightings/12112/details.
96
Specimens examined. – CANADA. ALBERTA: Edmonton, Patricia Ravine, 53.503638, -
113.593736, 2020, on bark at base of large, live Picea glauca, D. Haughland 2020-49 & A. Hood (hb.
Haughland); Edmonton, MacKenzie Ravine, 1976, on Betula bark, D.C. Lindsay s.n. (PMAE-B77.24.158);
Edmonton, grassy park next to Saskatchewan Drive bordering river valley, 53.513197, -113.53866, 2021,
on Picea twigs, D. Haughland 2021-34 (hb. Haughland).
Polycauliona polycarpa (Hoffm.) Frödén, Arup, & Søchting
(≡ Xanthoria polycarpa (Hoffm.) Th. Fr. ex Rieber) FIGURE 32 B.
Rare river valley and parkland epiphyte. Thalli form small, rounded, bright-orange cushions
typically less than the size of a quarter. Stipitate apothecia typically are common but are lacking in one of
the Edmonton collections. Laminal, immersed pycnidia present, conidia averaging 2.7 × 1.3 µm. No
vegetative propagules. Like P. candelaria, this species lacks rhizines, instead attaching with small peg-like
haptors. Unlike P. candelaria, the lobes tend to be short and rounded. Historically the much more common
Xanthomendoza hasseana was misidentified as this species in Alberta; they can be differentiated by the
well-developed white rhizines often visible around the edges of the lobes of X. hasseana. Edmonton
material is poorly developed, but the morphology and pycnidia measurements fit those in Lindblom (1997).
Chemistry: orange-pigmented parts K+ purple (vs. K- in Candelaria), all other spot tests negative, parietin,
fallacinal, emodin, teloschistin, and parietinic acid by TLC (Lindblom 2004b). Molecular support: a
variable species in need of phylogenetic work, no new sequences generated. Genus-level support provided
by Arup et al. (2013).
Specimens examined. – CANADA. ALBERTA: Edmonton, grassy park next to Saskatchewan
Drive bordering river valley, 53.513211, -113.538619, 2021, on Picea, D. Haughland 2021-28 (hb.
Haughland); Edmonton, Mill Creek ravine, 53.517222, -113.473889, 2020, on Picea glauca, D. Haughland
2020-115B & P. Williams (hb. Haughland).
Rusavskia elegans (Link) S. Y. Kondr. & Kärnefelt
(≡ Xanthoria elegans (Link) Th. Fr.) FIGURE 32 C.
River valley and parkland saxicole. More common in arid habitats, this dusky-orange species is
almost crustose in habit, closely attached to the substrate, and lacks rhizines. Thalli are composed of
narrow, convex, radiating lobes, typically with abundant apothecia centrally, and lacking isidia or soredia.
To discriminate it from Caloplaca, check that the lobes have an intact, white lower cortex, attached with
short peg-like hapters (often sparse). Typically, there is also space between the lobes where the substrate is
visible. The only record in Edmonton is from an online app, however recent informal observations suggest
it is common on headstones in cemeteries. Chemistry: orange-pigmented parts K+ purple (vs. K- in
Candelaria), all other spot tests negative, parietin, fallacinal, emodin, teloschistin, and parietinic acid by
TLC (Lindblom 2004b). Molecular support: well-supported at genus-level in existing phylogenetic
analyses (Arup et al. 2013), no new sequences generated.
Specimen observation. – CANADA. ALBERTA: Edmonton, Mill Creek Ravine South,
53.508699, -113.461606, 2019, on metal footbridge, T.L. Dueck (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/9806/details).
Vulpicida pinastri (Scop.) J.-E. Mattsson & M. J. Lai FIGURE 31 C.
River valley epiphyte. This common boreal species forms almost fluorescent yellow, ascending,
irregular, foliose rosettes, most commonly on Betula in river valley parks in Edmonton. The ruffled lobe
edges are lined with marginal, powdery soralia marginally. The color can vary to pale greenish-yellow in
shade forms. The rhizines are sparse, brownish-white, and develop from the whitish, wrinkled lower cortex.
Apothecia are not known from Alberta. Chemistry: all spot test negative, usnic acid, pinastric acid, zeorin,
and vulpinic acid by TLC (Brodo et al. 2001). Molecular support: the species is well-supported by
phylogenetic analyses in Saag et al. (2014) and is the sole soredia-forming clade within the genus, no new
sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Hawrelak Park, 53.52, -113.54, 2017, Betula papyrifera, D. Haughland et al. (unvouchered observation);
Edmonton, Rio Park, 53.505094, -113.595550, 2020, on Betula, L. Hjartarson (unvouchered observation:
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NatureLynx record https://naturelynx.ca/sightings/12101/details); Edmonton, MacKenzie Ravine, 1976, on
rotting log, D.C. Lindsay s.n. (PMAE-B77.24.41).
Xanthomendoza fallax (Hepp ex Arnold) Søchting, Kärnefelt & S. Y. Kondr.
(≡ Xanthoria fallax (Hepp ex Arnold) Arnold) FIGURE 32 D.
Ubiquitous and abundant epiphyte across all habitats. This species is characterized by yellow-
orange to dark orange appressed, foliose thalli with crescent-shaped marginal soralia developing greenish-
yellow powdery to grainy soredia. The soralia form "bird nest"-like shapes in a split between the upper and
lower cortices. Somewhat similar to X. fulva except for soralia development (submarginal to labriform in X.
fulva), X. fallax also has more abundant rhizines and typically wider lobes. Chemistry: orange parts K+
purple, all other spot tests negative, parietin, fallacinal, emodin, teloschistin and parietinic acid by TLC
(Lindblom 2004a). Molecular support: sequences in GenBank form a well-supported clade in preliminary
analyses (Haughland, unpub. data), no new sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Urban Monitoring Site 16, 53.468373, -113.59264, 2019, on trunk of Fraxinus pennsylvanica, M.
Villeneuve & M. Lewis s.n. [UoA-CC-73] (hb. Haughland); Edmonton, Urban Monitoring Site 46,
53.509321, -113.574933, 2019, on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n. [UoA-
CC-155] (hb. Haughland); Edmonton, Urban Monitoring Site 78, 53.547665, -113.502375, 2019, on trunk
of Tilia, D. Haughland & M. Cao s.n. [UoA-CC-159] (hb. Haughland); Edmonton, Urban Monitoring Site
111, 53.587075, -113.457605, 2019, on trunk of Fraxinus pennsylvanica, S. Toni & A. Hood s.n. [UoA-CC-
96] (hb. Haughland); Edmonton, Urban Monitoring Site 185E, 53.62906, -113.56883, 2019, on trunk of
Populus, D. Thauvette & M. Cao s.n. [UoA-CC-91] (hb. Haughland); Woodcroft, Air Quality Monitoring
Station, 53.563708, -113.563508, 2019, on trunk of Prunus virginiana, D. Haughland & A. Hood s.n.
[UoA-CC-107] (hb. Haughland); Edmonton, Sir Wilfrid Laurier Park, 53.50834, -113.560926, 2019, on
bark of large downed Populus, D. Haughland 2019-115 & P. Williams (hb. Haughland); Edmonton,
Government House, 53.542066, -113.545148, 2013, on mature Ulmus in lawn W of building, D.
Haughland 2013-01 (hb. Haughland); Edmonton, Henrietta Muir Edwards Park, near Accidental Beach,
53.538684, -113.470833, 2019, on downed wood, T.L. Dueck (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/7470/details); Edmonton, between Provincial Museum parking lot
and Wellington Crescent, 1976, on N sides of trunks of deciduous trees, D.C. Lindsay s.n. (PMAE-
B77.24.46); Edmonton, between Stony Plain Rd. and 100 Ave. at 148 St., 1976, on wooden footbridge,
D.C. Lindsay s.n. (PMAE-B77.24.66); Edmonton, Terwillegar Park, 1977, on wood, D.C. Lindsay s.n.
(PMAE-B77.24.81); Edmonton, Emily Murphy Park, 1976, on Betula bark, D.C. Lindsay s.n. (PMAE-
B77.24.87); Edmonton,Whitemud Park, 1976, on wood, D.C. Lindsay s.n. (PMAE-B77.24.109).
Xanthomendoza fulva (Hoffm.) Søchting, Kärnefelt & S. Y. Kondr.
(≡ Xanthoria fulva (Hoffm.) Poelt & Petut.) FIGURE 32 E.
Overlooked tableland epiphyte. A diminutive species forming thalli under 1 cm in diameter, it can
be confused with poorly developed X. fallax. Characterized by small, dark-red to orange thalli with rounded
or finely divided lobes that end in submarginal to labriform soralia. It often grows in small quantities with
the more abundant fallax. Given its air quality indicator value (Schulze et al. 2020), we predict this lichen
has been overlooked and is more abundant on boulevard trees in Edmonton than our data suggests.
Chemistry: orange parts K+ purple, all other spot tests negative, parietin, fallacinal, emodin, teloschistin
and parietinic acid by TLC (Lindblom 2004a). Molecular support: sequences in GenBank form a well-
supported clade in preliminary analyses (Haughland, unpub. data), no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 4, 53.443624,
-113.461366, 2019, on trunk of Ulmus americana, D. Haughland & A. Stordock s.n. [UoA-CC-141] (hb.
Haughland); Edmonton, Urban Monitoring Site 7, 53.452098, -113.573622, 2019, on trunk of Ulmus
americana, D. Haughland & L. Hjartarson s.n. [UoA-CC-40,143] (hb. Haughland); Edmonton, Urban
Monitoring Site 83, 53.559591, -113.642062, 2019, on trunk of Ulmus americana, D. Haughland & L.
Hjartarson s.n. [UoA-CC-34,160] (hb. Haughland).
98
Figure 33. Orange foliose lichens of Edmonton, plate 2 of 2: introduced Xanthoria parietina. A, Colonies
on planted suburban Hydrangea tree, Garvey https://naturelynx.ca/sightings/15817/details. B, Colonies on
horticultural Malus trees imported to Alberta from a greenhouse in Surrey, British Columbia, Haughland
unvouchered observation.
Xanthomendoza hasseana (Räsänen) Søchting, Kärnefelt & S. Y. Kondr.
(≡ Xanthoria hasseana Räsänen) FIGURE 32 F.
Occasional tableland, parkland, and river valley and ravine system epiphyte. Forming orange
rosettes, this species typically has at least some apothecia and laminal, slightly protuberant orange pycnidia.
Rhizines often are visible around the edges of the overlapping, convex lobes. Historically this species was
misidentified as Xanthoria polycarpa in Alberta, a species that lacks rhizines and has a more compact
growth habit (see Polycauliona polycarpa entry above). Xanthomendoza hasseana thalli are diminutive
within Edmonton and easily overlooked and misidentified as X. fallax, which commonly is fertile in the
city. It differs from X. fallax in lacking soredia and the more prominent contrasting pycnidia.
Xanthomendoza montana (L. Lindblom) Søchting, Kärnefelt & S. Y. Kondr. is a very similar species that
appears to be less common in Alberta, and is differentiated from X. hasseana with difficulty by spore and
septum size (Lindblom 1997; measurements from Brodo 2016: X. montana spores (11.5–)12.5–15.5 × 5–
7.5 μm, septum about one-third the length of the spore or less, 1.5–4 μm wide vs. X. hasseana spores 15–18
× 7.0–8.0(–9.5), septum more than one-third the length of the spore, 4–8.5 μm wide). In our experience,
spores often show intermediate values or the spore size fits one species while the septum width fits the
other. Chemistry: orange parts K+ purple, all other spot tests negative, parietin, fallacinal, emodin,
teloschistin and parietinic acid by TLC (Lindblom 2004a). Molecular support: no new sequences generated,
in preliminary analyses (Haughland, unpub. data) sequences in GenBank form multiple well-supported
clades, some intermixed with X. montana. Species-level phylogenetic analyses are needed.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Urban Monitoring Site 33, 53.479311, -113.413258, 2019, on trunk of Tilia, D. Thauvette et al. s.n. [UoA-
CC-147] (hb. Haughland); Edmonton, Urban Monitoring Site 85, 53.560830, -113.567591, 2019, on trunk
of Fraxinus pennsylvanica, D. Haughland & A. Hood s.n. [UoA-CC-162] (hb. Haughland); Edmonton,
Urban Monitoring Site 0E, 53.442169, -113.584786, 2019, on trunk of Ulmus americana, D. Haughland &
L. Hjartarson s.n. [UoA-CC-64] (hb. Haughland); Edmonton, Urban Monitoring Site 175E, 53.616551, -
113.562121, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland s.n. [UoA-CC-24] (hb. Haughland);
Edmonton, Urban Monitoring Site 34E, 53.471215, -113.371731, 2019, on trunk of Fraxinus
pennsylvanica, D. Thauvette & S. Toni s.n. [UoA-CC-148] (hb. Haughland); Edmonton, South Air Quality
Monitoring Station, 53.501931, -113.523499, 2019, on trunk of Fraxinus, D. Thauvette & J. Birch s.n.
[UoA-CC-189] (hb. Haughland); Edmonton, Patricia Ravine, 53.503513, -113.59324, 2020, epiphytic, L.
99
Hjartarson (unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/12112/details);
Edmonton, approx. 180 St. and 99 Ave., 1976, on twigs and dead branches, D.C. Lindsay s.n (PMAE-
B77.24.176); Edmonton, Terwillegar Park, 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.177).
*Xanthoria parietina (L.) Th. Fr. FIGURE 33.
Rare introduced tableland epiphyte. This is a distinctive yellow-orange foliose lichen, with thin,
broad, concave lobes that resemble petals, with abundant apothecia centrally. No vegetative propagules.
The only lichen we are confident was introduced to Alberta, it appears to be limited to ornamental
horticultural trees and shrubs originally grown on the west coast and shipped to Alberta for sale (see also
Brodo et al. 2021a). We previously examined nature app records for proof of this species in Alberta, but
reports were misidentifications. Here we provide two verified observations, one of multiple thalli on a
Hydrangea tree that was planted approximately ten years ago, and a second, recent observation of
additional thalli on newly imported greenhouse trees and shrubs from Abbotsford, British Columbia.
Chemistry: yellow-orange parts K+ purple, all other spot tests negative, parietin, fallacinal, emodin,
teloschistin and parietinic acid by TLC (Lindblom 2004a). Molecular support: no new sequences generated,
in preliminary analyses (Haughland, unpub. data) sequences in GenBank form multiple well-supported
clades. The biogeography and genetic structure within X. parietina potentially could be a fascinating study
of range expansion in a “weedy” lichen.
Specimens examined. – CANADA. ALBERTA: Edmonton, 53.504211, -113.66915, 2021, on
ornamental Hydrangea tree, H. Davidson s.n. (hb. Spribille, NatureLynx record
https://naturelynx.ca/sightings/15817); Edmonton, Spruce Avenue neighbourhood, 53.563562, -
113.498138, 2020, on ornamental Syringia shrub, D. Haughland s.n.
GROUP 9. WHITE AND GREY FOLIOSE LICHENS
Nine species. Key literature: Brodo et al. 2013; Brodo 2016; Esslinger 2016a, 2016b, 2016c;
Esslinger et al. 2020; Goward et al. 1994; Lendemer 2009; Lendemer & Hodkinson 2010.
1a. Lobes hollow tubes, lacking cilia or rhizines; soralia terminal, hood-shaped ........ Hypogymnia physodes
1b. Lobes solid and flattened, typically with cilia or rhizines; soralia variable ............................................... 2
2a. Lobes >3 mm wide .................................................................................................................................... 3
2b. Lobes 0.5-2.5 mm wide ............................................................................................................................. 4
3a. Upper surface smooth to wrinkled, lobe edges rounded, pseudocyphellae laminal, punctiform to slightly
elongate, soralia marginal to laminal; lower surface tan, with short, pale, simple rhizines ...............................
............................................................................................................................................. Punctelia caseana
3b. Upper surface with hammered appearance, lobe edges angular, pseudocyphellae forming a network
along laminal ridges and lobe margin, developing into linear soralia; lower surface black with black
rhizines that become ‘bottle-brush-like’ with squarrose branching ....................................... Parmelia sulcata
4a. Upper cortex with a subtle combed or flowing appearance; lower cortex lacking; rhizines black, with
‘bottle-brush’ squarrose branching .............................................................................. Heterodermia japonica
4b. Upper cortex with smooth, matte, shiny or hammered appearance; lower cortex present, white to pale
brown to black; rhizines variable, mostly simple or sparsely branched, concolorous with lower cortex ........ 5
5a. Lower cortex dark at least centrally; rhizines typically bristling around lobes, black with white tips;
soralia laminal, rounded to irregular in outline; apothecia occasional, typically with cilia on thalline
exciple; upper cortex K- .......................................................................................... Phaeophyscia orbicularis
5b. Lower cortex pale to tan throughout; rhizines sparse, white to pale brown; soralia present or absent,
when present variable, apothecia present or absent but when present lacking cilia; upper cortex K+ yellow
(atranorin) ....................................................................................................................................... 6 (Physcia)
6a. Lobes ascending, with long marginal cilia; lobe tips with helmet-shaped inflated soralia; thallus highly
variable from short rounded lobes to sparse elongate lobes ............................................. Physcia adscendens
6b. Lobes appressed to substrate, lacking marginal cilia; esorediate or with marginal, elongate soralia ........ 7
100
7a. Soredia abundant in marginal soralia, apothecia common with sorediate thalline margin
....................................................................................................................................... Physcia aff. dimidiata
7b. Esorediate, apothecia typically abundant and with smooth, esorediate thalline margin ............................ 8
8a. Lobes convex, clasping substrate; apothecia slightly elevated above thallus and with relatively thick
thalline rim; upper cortex white to pale grey, lacking maculae or maculae indistinct; medulla K- ...................
......................................................................................................................................... Physcia aff. stellaris
8b. Lobes flattened to slightly concave; apothecia attached closely to thallus with relatively thin thalline
rim; upper cortex often maculate and blueish grey; medulla K+ yellow ................. 9 (Physcia aipolia group)
9a. Apothecia in center of thallus often relatively large with smaller apothecia towards mid-thallus; lobes
relatively wide (to >1 mm) with little substrate visible between lobes .................................... Physcia aipolia
9b. Apothecia typically similar in size from center to edge of thallus, smaller and spread out to lobe tips;
lobes narrow (≤0.8 mm) and substrate visible between lobes
................................................................................ Physcia alnophila [see discussion under Physcia aipolia]
*Heterodermia japonica (M. Satô) Swinscow & Krog
(≡ Polyblastidium japonicum (M. Satô) Kalb) FIGURE 34.
We have not adopted the taxonomy of Mongkolsuk et al. (2015) while we wait on the work of T.
Esslinger and S. Leavitt on Physciaceae in North America, and collaborative molecular work with T.
Spribille. Rare river valley and ravine system epiphyte. This species was first detected in Alberta by the
ABMI and confirmed by J. Lendemer for the senior author in 2013, however, we have not formally
reported its presence until now. In Canada, this species has also been recorded from British Columbia and
Ontario (CNALH 2020). Herbarium collections show this species has been overlooked in Alberta for
decades, historically misidentified as Physcia tenella (Scop.) DC. or Heterodermia speciosa (Wulfen)
Trevisan (Haughland, unpubl.). It appears to be an occasional northern forest epiphyte. Lobes long, plane,
appressed, epruinose, upper cortex prosoplectenchymatous, ecorticate lower surface. Soralia marginal.
Rhizines marginal to terminal, dark and squarrose. A single historical specimen was located in Edmonton;
recent searches in the area failed to find extant thalli, but contemporary specimens were observed
immediately north of the city. Chemistry: K+ yellow, PD+ faint yellow, all other spot tests negative.
Secondary metabolites detected by TLC: atranorin, zeorin, unknown terpenes. Molecular support: no new
sequences generated, good species-level support for sequences from Costa Rica (Lücking et al. 2008).
Specimens examined. – CANADA. ALBERTA: Edmonton, MacKenzie Ravine, 1976, on bark,
D.C. Lindsay s.n. (PMAE-B77.24.164); St. Albert, Riverlot 56 Natural Area, 53.6585, -113.5849, 2020, on
Populus balsamifera, D. Haughland 2020-112 (hb. Haughland, NatureLynx record
https://naturelynx.ca/sightings/14920/details).
Hypogymnia physodes (L.) Nyl. FIGURE 35 A.
Occasional river valley and ravine system epiphyte in Edmonton. While relatively rare in the city,
it is an extremely common epiphyte elsewhere in Alberta. Hollow-lobed, highly variable, white to greenish
on upper half, with a blackened lower cortex lacking rhizines. Lobes can be long or short, appressed or
ascending, usually 1–2.5(–5) mm across. Underside of lobe tips bursting into hooded soralia. One of the
more commonly used species for biomonitoring. Chemistry: cortex K+ yellow, medulla K+ slow red to
dingy brown, PD+ orange. Secondary metabolites detected by TLC: atranorin, physodic acid, physodalic
acid, protocetraric acid, 3-hydroxyphysodic acid. Molecular support: species is monophyletic in
Miadlikowska et al. (2011), no new sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Patricia Ravine, 53.503216, -113.592645, 2020, on Prunus stems, D. Haughland & A. Hood (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/14352/details); Edmonton, River Valley,
Oleskiw, 53.499071, -113.59265, 2020, epiphytic, samanthapedersen (unvouchered observation:
iNaturalist record https://www.inaturalist.org/observations/42400752); Edmonton, MacKenzie Ravine,
1976, on rotting trunk, D.C. Lindsay s.n (PMAE-B77.24.42).
101
Figure 34. White and grey foliose lichens of Edmonton, plate 1 of 2: Heterodermia japonica (collections
imaged are from outside of Edmonton, citations provided here). A, Thallus, ca. 26 km NE of Marten Beach,
ABMI Site 720, 55.68, -114.67, 2003, on hard log, E. Bainbridge & J. Bluetchen s.n. [ABMI Lichen #
528428]. B-C, ca. 51 km NE of Sandy Lake, ABMI Site 664, 55.85, -112.61, 2015, on log, S. Venskaitis
s.n. [ABMI Lichen # 626478]. B, Lower surface showing lack of cortex and squarrose rhizines. C, Upper
surface with marginal soralia and prosoplectenchymatous upper cortex.
Parmelia sulcata Taylor FIGURE 35 B.
River valley, parkland, and rare tableland epiphyte and xylicole. Perhaps the most abundant
foliose epiphyte in Alberta, this species is surprisingly sparse in Edmonton outside of river valley parks.
Thallus blue-grey with angled lobes, often browned at tips in exposed sites. Lobes with a network of sharp
ridges and depressions and whitish pseudocyphellae, developing into powdery soredia on lobe margins and
ridges. Rhizines typically densely squarrose-branched, but young rhizines often simple. Black lower cortex.
Apothecia rare. Chemistry: cortex K+ yellow, PD-, medulla K+ red, PD+ orange, other spot tests negative.
Substrances detected by TLC: atranorin, salazinic acid, ±consalazinic acid. Molecular support: strong
species-level support (Molina et al. 2017), no new sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Mill
Creek Ravine, 2020, 53.507682, -113.465167, gpohl (unvouchered observation: iNaturalist record
102
https://www.inaturalist.org/observations/49307449); Edmonton, 10345 144 St. NW, 53.544996, -
113.567651, 2017, epiphytic, jwyattpsd70 (unvouchered observation: iNaturalist record https://www.
inaturalist.org/observations/7527077); Edmonton, Mill Creek Ravine, 53.527788, -113.479163, 2020,
tawneedupuis (unvouchered observation: iNaturalist record https://www.inaturalist.org/obser
vations/56905704); Edmonton, Whitemud Park, 53.49903, -113.560691, 2020, on tree base, hanna1025
(unvouchered observation: iNaturalist record https://www.inaturalist.org/observations/59075224);
Edmonton, MacKenzie Ravine, 1976, on tree, D.C. Lindsay s.n. (PMAE-B77.24.32); Edmonton,
McKinnon Ravine, 1976, on Populus bark, D.C. Lindsay s.n. (PMAE-B77.24.56); Edmonton, approx. 1
mile W of 95 Ave. and 170 St. intersection, 1975, on Populus tremuloides, D.C. Lindsay s.n (PMAE-
B77.24.62); Edmonton, between Stony Plain Rd. and 100 Ave. at 148 St., 1976, on wooden footbridge,
D.C. Lindsay s.n. (PMAE-B77.24.63); Edmonton, approx. 180 St. and 99 Ave., 1976, on twigs and dead
branches, D.C. Lindsay s.n. (PMAE-B77.24.69); Edmonton, Terwillegar Park, 1977, on wood, D.C.
Lindsay s.n. (PMAE-B77.24.77); Edmonton, Near Northland Sandpit, 2 mi W and 1 mi S of 170 St. and 79
Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.103); Edmonton, Terwillegar Footbridge, 53.4797,
-113.594315, 2021, on Picea snag, D. Haughland 2021-2B (hb. Haughland); Edmonton, Hawrelak Trail
off-leash park, 53.522361, -113.54378, 2020, on decorticate Picea twigs, D. Haughland 2020-25 (hb.
Haughland).
Physcia adscendens (Fr.) H. Olivier FIGURE 35 D.
Common and variable epiphyte across all treed habitats in Alberta, including Edmonton. Thalli to
2 cm in diameter, mostly irregular, with ascending lobes attaching to substrate centrally. Lobes widest at
tips due to abundant helmet-shaped soralia. Marginal cilia abundant, darkening distally. Upper cortex white
to grey, sometimes maculate, never pruinose, lower cortex typically white. Chemistry: cortex K+ yellow,
all other spot tests negative. Secondary metabolites detected by TLC: atranorin. Molecular support: work is
required given the morphological diversity evidenced even within Alberta, samples submitted to T.
Esslinger and S. Leavitt (unpublished data).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Urban Monitoring Site 11, 2019, on trunk of Fraxinus pennsylvanica, 53.450690, -113.474695, D.
Thauvette & J. Wasyliw s.n. [UoA-CC-101] (hb. Haughland); Edmonton, Urban Monitoring Site 39,
53.494317, -113.525188, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & A. Stordoc s.n.
[UoA-CC-154] (hb. Haughland); Edmonton, Urban Monitoring Site 76, 53.547844, -113.549140, 2019, on
trunk of Ulmus americana, D. Haughland & A. Hood s.n. [UoA-CC-3,53] (hb. Haughland); Edmonton,
Urban Monitoring Site 102, 53.574582, -113.435207, 2019, on trunk of Ulmus americana, D. Royko & R.
Fielder s.n. [UoA-CC-170] (hb. Haughland); Edmonton, Urban Monitoring Site 174E, 53.612602, -
113.589391, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland s.n. [UoA-CC-22] (hb. Haughland);
Edmonton, Beverly Air Quality Monitoring Station, 53.566860, -113.397464, 2019, on trunk of Ulmus
americana, D. Thauvette & J. Birch s.n. [UoA-CC-13,26,138,188] (hb. Haughland); Edmonton, Buena
Vista Park, 53.518208, -113.548277, 2020, on tree-form Salix, D. Haughland (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/12611/details); Edmonton, between Provincial Museum
parking lot and Wellington Crescent, 1975, on N sides of trunks of deciduous trees, D.C. Lindsay s.n.
(PMAE-B77.24.48); Edmonton, approx. 180 St. and 99 Ave., 1976, on twigs and dead branches, D.C.
Lindsay s.n. (PMAE-B77.24.73); Edmonton, Terwillegar Park, 1977, on wood, D.C. Lindsay s.n. (PMAE-
B77.24.82); Edmonton, Emily Murphy Park, 1976, on Betula bark, D.C. Lindsay s.n. (PMAE-B77.24.89);
Edmonton, MacKenzie Ravine, 1976, on bark, D.C. Lindsay s.n. (PMAE-B77.24.157).
Physcia aipolia (Ehrh. ex Humb.) Fürnr. group FIGURE 35 E.
River valley, parkland, and tableland epiphyte. This species is rarer than the somewhat similar P.
aff. stellaris in Edmonton. Here we use “group” as Edmonton likely has both P. aipolia and P. alnophila
(Vain.) Loht., Moberg, Myllys & Tehler. Broader efforts through the ABMI to distinguish these two
species based on morphology and TLC (e.g., Brodo et al. 2013) left us doubting our ability to accurately
discriminate them (Haughland, unpublished data) so at present we prefer to group them. The key provides
some field traits other authors have found useful to separate P. alnophila versus P. aipolia, however, TLC
can aid in a definitive identification (Brodo et al. 2013). Edmonton material: thallus orbicular to irregular,
up to 5 cm in diameter, whitish-grey to dark grey, typically maculate. Lobes flattened to slightly concave
103
(especially at the tips, vs. convex and clasping in P. aff. stellaris). Lacking soredia or isidia, apothecia
common, appressed on thallus (vs. stipitate in P. aff. stellaris), thalline margin relatively thin (vs. relatively
thick in P. aff. stellaris), disk often pruinose (vs. typically epruinose in P. aff. stellaris). Chemistry: cortex
and medulla K+ yellow (vs. medulla K- in P. aff. stellaris), all other spot tests negative. Secondary
metabolites detected by TLC: atranorin, zeorin, multiple unknown fatty acids and terpenes. Molecular
support: work is required, samples submitted to T. Esslinger and S. Leavitt (unpublished data).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 9, 53.453715, -113.527142, 2019, on trunk of Fraxinus, D. Haughland & A. Stordock s.n. [UoA-CC-
144] (hb. Haughland); Edmonton, Urban Monitoring Site 46, 53.509321, -113.574933, 2019, on trunk of
Ulmus americana, D. Haughland & L. Hjartarson s.n. [UoA-CC-155] (hb. Haughland); Edmonton, Urban
Monitoring Site 86, 53.561118, -113.547994, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland &
A. Hood s.n. [UoA-CC-163] (hb. Haughland); Edmonton, Urban Monitoring Site 100, 53.574491, -
113.479536, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-CC-169] (hb.
Haughland); Edmonton. Urban Monitoring Site 175E, 53.616594, -113.562214,2019, on trunk of Fraxinus
pennsylvanica, D. Haughland s.n. [UoA-CC-25] (hb. Haughland); Ardrossan, Air Quality Monitoring
Station, 53.554823, -113.143428, 2019, on trunk of Populus balsamifera, D. Thauvette & J. Birch s.n.
[UoA-CC-187] (hb. Haughland); Edmonton, Patricia Ravine, 53.504611, -113.593583, 2020, on upper
branches of downed Populus balsamifera, D. Haughland 2020-55 & A. Hood (hb. Haughland); Edmonton,
between Stony Plain Rd. and 100 Ave. at 148 St., 1976, on supports of wooden footbridge and on bark,
D.C. Lindsay s.n. (PMAE-B77.24.53); Edmonton, MacKenzie Ravine, 1976, on bark, D.C. Lindsay s.n.
(PMAE-B77.24.154); Edmonton, Whitemud Park, 1976, on wood, D.C. Lindsay s.n. (PMAE- as minor
component in B77.24.125); Edmonton, grassy park next to Saskatchewan Drive bordering river valley,
53.513211, -113.538619, 2021, on Picea twigs, D. Haughland 2021-29 (hb. Haughland).
***Physcia aff. dimidiata (Arnold) Nyl. FIGURE 35 F.
River valley epiphyte, occasional in parkland and tableland habitats. In contrast to the Sonoran
region where P. dimidiata occurs on northern-exposed rock (Moberg 2002), in Alberta these specimens are
epiphytic, occasionally forming luxurious colonies on conifer twigs in riparian zones. Edmonton material:
thallus loosely attached, grey-white with a dense crystalline pruina, and lobes with crenulate margins
bearing granular soredia. Commonly apotheciate, with soralia forming on eroded thalline apothecial
margin. Chemistry: cortex K+ yellow, medulla K-, all other spot tests negative, secondary metabolites not
investigated. Molecular support: in Alberta, this species is not monophyletic and the material does not
group with P. dimidiata from the southwestern United States (T. Esslinger, pers. comm.). Molecular work
is ongoing as part of a comprehensive treatment of Physciaceae by T. Esslinger and S. Leavitt, but it is
clear this material does not cluster with Physcia dimidiata s.s.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 7, 53.452098, -113.573622, 2019, on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n.
[UoA-CC-143] (hb. Haughland); Edmonton, Urban Monitoring Site 38, 53.498347, -113.546635, 2019, on
trunk of Ulmus, D. Haughland & A. Stordoc s.n. [UoACC-152] (hb. Haughland); Edmonton, Urban
Monitoring Site 39, 53.494317, -113.525188, 2019, on trunk of Fraxinus pennsylvanica, D. Haughland &
A. Stordoc s.n. [UoA-CC-154] (hb. Haughland); Edmonton, Urban Monitoring Site 46, 53.509380, -
113.574106, 2019, on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n. [UoA-CC-131] (hb.
Haughland); Edmonton, Urban Monitoring Site 69, 53.534195, 113.525517, 2019, on trunk of Populus
balsamifera, D. Haughland & S. Toni s.n. [UoA-CC-126] (hb. Haughland); Edmonton, near Whitemud
Park, S bank of North Saskatchewan River, 53.5058, -113.5551, 2017, epiphytic on branches and bark of
young Picea glauca, D. Haughland 2017-235a & P. Williams (hb. Haughland), D. Haughland 2017-235b
& P. Williams (hb. Haughland); Edmonton, Hawrelak Trail off-leash park, 53.51807, -113.5401, 2017, on
Picea glauca branches & twigs, coating lower dead branches, D. Haughland 2017-233 & P. Williams (hb.
Haughland); Edmonton, Hawrelak Trail off-leash park, 53.5134, -113.5395, 2017, on Picea glauca twigs &
branches, coating lower dead branches, D. Haughland 2017-234 & P. Williams (hb. Haughland);
Edmonton, MacKenzie Ravine, 1976, on bark, D.C. Lindsay, s.n. (PMAE-B77.24.166); Edmonton, grassy
park next to Saskatchewan Drive bordering river valley, 53.513211, -113.538619, 2021, on Picea twigs, D.
Haughland 2021-27 (hb. Haughland); Edmonton, Hawrelak Trail off-leash park, 53.522361, -113.54378,
2020, on decorticate Picea twigs, D. Haughland 2020-25 (hb. Haughland).
104
Figure 35. White and grey foliose lichens of Edmonton, plate 2 of 2. A, Hypogymnia physodes growing
with Parmelia sulcata (right), Haughland https://naturelynx.ca/sightings/14287/details. B, Parmelia
sulcata, Haughland unvouchered observation. C, Phaeophyscia orbicularis, Hjartarson https://
naturelynx.ca/sightings/11762/details. D, Physcia adscendens, Haughland https://naturelynx.ca
/sightings/12611/details. E, Physcia aipolia group, Haughland unvouchered observation. F, Physcia aff.
dimidiata, Haughland unvouchered observation. G, Physcia aff. stellaris, Haughland
https://naturelynx.ca/sightings/12612/details. H, Punctelia caseana, Haughland unvouchered observation.
105
***Physcia aff. stellaris (L.) Nyl. FIGURE 35 G.
Common tableland and parkland epiphyte, occasional in the river valley. Esslinger et al. (2020)
suggest that this species in the strict sense is not present in North America, instead, we have a complex of
similar but largely undescribed species. Until the complex is resolved, we lump these under P. aff. stellaris.
Material included here shares the following traits: thallus orbicular to irregular, to 5 cm in diameter. Thalli
varying from narrow lobed with apothecia clustered centrally to thalli consisting largely of clustered
apothecia. Lobes grey to white, upper surface may be maculate, pruinose or neither, typically convex and
slightly elevated and clasping the substrate. Apothecia common, slightly stipitate with a relatively thick
thalline margin, disk black and seldom pruinose. Includes wide-lobed, draping form with elevated
apothecia. No vegetative propagules. See P. aipolia group entry for comparison. Chemistry: cortex K+
yellow, medulla K-, all other spot tests negative. Secondary metabolites detected by TLC: atranorin.
Molecular support: work is required, samples submitted to T. Esslinger and S. Leavitt (unpublished data).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Urban Monitoring Site 14, 53.447673, -113.412609, 2019, on trunk of Ulmus americana, D. Royko & D.
Fielder s.n. [UoA-CC-146] (hb. Haughland); Edmonton, Urban Monitoring Site 33, 53.479375, -
113.412798, 2019, on trunk of Tilia, D. Thauvette et al. s.n. [UoA-CC-46] (hb. Haughland); Edmonton,
Urban Monitoring Site 119, 53.602014, -113.458033, 2019, on trunk of Fraxinus pennsylvanica, D. Royko
& D. Fielder s.n. [UoA-CC-176] (hb. Haughland); Edmonton, Urban Monitoring Site 150E, 53.590398, -
113.588890, 2019, on trunk of Ulmus americana, D. Haughland & L. Hjartarson s.n. [UoA-CC-134] (hb.
Haughland); Edmonton, South Air Quality Monitoring Station, 53.503437, -113.523856, 2019, on trunk of
Fraxinus, D. Thauvette & J. Birch s.n. [UoA-CC-4] (hb. Haughland); Edmonton, Buena Vista Park,
53.518211, -113.548263, 2020, on tree-form Salix, D. Haughland (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/12612/details); Edmonton, between Provincial Museum parking lot
and Wellington Crescent, 1975, on N sides of trunks of deciduous trees, D.C. Lindsay s.n. (PMAE-
B77.24.49); Edmonton, approx. 1 mile W of 95 Ave. and 170 St. intersection, 1975, on Populus
tremuloides, D.C. Lindsay s.n. (PMAE-B77.24.64); Edmonton, Emily Murphy Park, 1976, on Betula bark,
D.C. Lindsay s.n. (PMAE-B77.24.86); Edmonton, Whitemud Park, 1976, on wood, D.C. Lindsay s.n.
(PMAE-B77.24.125); Edmonton, along Saskatchewan Drive, near the Biological Sciences Building of the
University of Alberta, 1976, on bark (cf. Salix), D.C. Lindsay s.n. (PMAE-B77.24.179 ); Edmonton, 98
Ave. and 154 St. intersection, 1976, on bark, D.C. Lindsay s.n. (PMAE-B77.24.190); Edmonton,
McKinnon Ravine, 1976, on Populus bark, D.C. Lindsay s.n. (PMAE-B77.24.59); Edmonton, approx. 180
St. and 99 Ave., 1976, on twigs and dead branches, D.C. Lindsay s.n. (PMAE-B77.24.72).
Punctelia caseana Lendemer & Hodkinson FIGURE 35 H.
Occasional river valley epiphyte. Historically our material was called Punctelia subrudecta (Nyl.)
Krog in Alberta, but Lendemer & Hodkinson (2010) clarified that species and its North American
distribution. Alberta material was cited as P. jeckeri (Roum.) Kalb based in part on the presence of pruina
on the lobe tips (Lendemer & Hodkinson 2010; Elk Island National Park 1961, G.W. Scotter 657 [CANL],
not examined here). Our analyses show that material in Alberta is instead P. caseana, calling into question
the utility of pruina in distinguishing these species, particularly in base-rich/alkaline environments like
much of Alberta. Recognized by up-to-palm-sized thalli, with wide, rounded, grey lobes with primarily
laminal soralia producing coarse granular soredia that often aggregate. The lobe tips vary from epruinose to
sparsely pruinose, lobes are adnate except at the distal edges, and the lower surface is beige to pale brown
with short, simple, “buzz-cut”-like rhizines. Can be confused with Flavopunctelia that are low in usnic
acid, in which case examining the lower cortex (black in Flavopunctelia) or chemistry (cortex KC+ yellow
in Flavopunctelia) will distinguish them. Chemistry: cortex K+ yellow, medulla C+ red, KC+ red, all other
spot tests negative. Secondary metabolites detected by TLC: lecanoric acid, atranorin (trace). Molecular
support: high at genus and species level. Analyses to date, as well as those conducted herein suggest
relatively high support for P. caseana and reciprocal monophyly with P. jeckeri (Alors et al. 2016,
Lendemer & Hodkinson 2010; Fig. 9 herein).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Patricia Ravine, 53.503216, -113.592645, 2020, on Prunus stems, D. Haughland & A. Hood s.n.
(unvouchered observation); Edmonton, Larch Sanctuary, 53.453522, -113.547402, 2020, epiphytic, D.
Haughland (unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/10688/details);
106
Edmonton, Hawrelak Park, 53.51665, -113.538811, 2020, on Betula papyrifera, D. Haughland 2020-111
(hb. Haughland).
GROUP 10. BROWN AND GREEN FOLIOSE LICHENS
Eleven species. Key literature: Breuss 2002; Brodo 2016; Esslinger 1977; Esslinger et al. 2017. A
group of often overlooked species, typically camouflaging well with the bark they commonly grow on.
With practise most species can be identified in the field.
1a. Squamulose; esorediate; with embedded perithecia .................................... Endocarpon aff. unifoliatum
1b. Foliose; mostly sorediate and/or isidiate; if fertile with apothecia not perithecia ..................................... 2
2a. Lobes dark reddish to greenish brown, often shiny, typically appressed; lower cortex and rhizines
similar in color and sheen to upper cortex or darkening; rhizines simple .....3 (Melanelixia & Melanohalea)
2b. Lobes dull green, brown or grey; lower cortex black at least centrally; rhizines branching to densely
squarrose .......................................................................................................................................................... 6
3a. Esorediate; isidia, lobules and/or apothecia present .................................................................................. 4
3b. Sorediate; may also have very small granular isidia or sparse apothecia .................................................. 5
4a. Isidiate, isidia abundant, with a greasy luster, becoming large, hollow, and lobulate; apothecia rare;
pycnidia not observed; in riparian and tableland habitats in Edmonton ................ Melanohalea exasperatula
4b. Lacking isidia but may develop lobules; apothecia common; pycnidia abundant, laminal, immersed,
visible as black dots on upper surface; thus far known only from outskirts of Edmonton and not treated
further here ....................................................................................... Melanohalea subolivacea (see Fig. 37D)
5a. Soralia marginal, labriform, white; lacking isidia .................................................. Melanelixia albertana
5b. Soralia central, laminal, golden-green; with tiny cylindrical isidia ..................... Melanelixia subaurifera
6a. Upper surface lacking pruina, lower surface pale to darkening centrally; rhizines often bristling around
lobes, simple, black with white tips, but variable ................................................................. 7 (Phaeophyscia)
6b. Upper surface typically pruinose around margins; lower cortex dark black to brown; rhizines dense,
black and ‘bottle-brush’ squarrose ............................................................................................. 9 (Physconia)
7a. Lower cortex pale througout; lobe tips with blastidia and tiny hyaline hairs; thalli tiny, often
camouflaged against bark and easily overlooked .......................................................Phaeophyscia nigricans
7b. Lower cortex darkening at least centrally; lobes with terminal granular soredia or with laminal, rounded
soralia; thalli larger, more easily observed ...................................................................................................... 7
8a. Soralia marginal, with granular isidioid soredia/isidia with often conspicuous pale hairs; rare in
Edmonton in humid habitats ....................................................................................... Phaeophyscia kairamoi
8b. Soralia primarily laminal and sub-marginal, rounded in outline, the soredia mostly finely granular and
lacking hairs; ubiquitous .......................................................................................... Phaeophyscia orbicularis
9a. Soralia at end of lobes, labriform and discrete ........................................................................................ 10
9b. Soralia along edges of lobes, often continuous especially towards the center of the thallus, linear ........ 11
10a. Form ranges from rosettes to scattered lobes; lower surface typically pale and ecorticate; medullary
hyphae visible on lower surface darkening to form fine brown/black striations near lobe tips and a dark,
dull lower cortex forming towards the thallus center .................................................... Physconia perisidiosa
10b. Typically forming rosettes; lower surface abruptly darkening with a well-developed cortex past
soralia, obscuring medulla ................................................................................................... Physconia labrata
11a. Medulla and soralia white, K-, KC-............................................................................. Physconia detersa
11b. Medulla and soralia yellowish, K+ yellow, KC+ yellow to orange .................. Physconia enteroxantha
107
Figure 36. Endocarpon aff. unifoliatum, growing on indoor-outdoor carpeting in Edmonton, Haughland
2020-98. A, Squamules. B, Cross-section of perithecium. C, Squamule cross-section.
**Endocarpon aff. unifoliatum T. Zhang, X. L. Wei & J. C. Wei FIGURE 36.
Anthropogenic tableland habitat. The Verrucariaceae is a challenging family in Alberta, with
many of the collections the senior author has received through the ABMI not fitting existing North
American keys and species descriptions. This sample was no different. Molecular and morphological data
suggest that this material constitutes the first North American record of a recently described species from
China, Endocarpon unifoliatum (Zhang et al. 2017). Edmonton material: adnate grey-brown squamules
from 2–7 mm diameter on their longest axis, with ascending, darkened margins at the very edges of the flat
to concave, lobulate thalli. The squamules do not form imbricate colonies, covering the substrate in a
108
mostly single layer. The squamules are 170–250 µm thick, thickest centrally, the upper cortex of
paraplectenchymatous cells in vertical columns, 12.5–30.0 µm thick, penetrating the algal layer in places,
and overlain with a very thin epinecral layer that is best visualized under polarized light. No other
polarizing crystals were found in cross-section. The algal layer is irregular on both the top and bottom, and
formed of vertical columns of algae. The medulla consists of cream to pale yellow
subparaplectenchymatous to loosely interwoven fibrous hyphae, with some swollen and some cylindrical
cells. The lower cortex is formed of paraplectenchymatous cells, mostly dark brown to black except where
the squamules are lifted off the surface (at the edges and in some places in ripples in the middle of the
squamules). Hyaline to dark rhizohyphae 3–4 µm in diameter form irregular wefts on the lower surface,
and may form actual rhizines; the substrate (thin, tightly woven outdoor carpet glued on concrete) may be
impeding their development. Abundant laminal, immersed perithecia are present, forming slightly darkened
extruding bumps on the upper surface, and protrusions through the lower surface. Perithecia 150–170 µm
wide and high, pyriform, with abundant paraphyses and a black exciple. Asci are 75–80 × 15 µm,
bisporous, with hyaline, muriform spores measuring 30–34 × 13–15 µm. The perithecia house small
globose algae, 1–2 celled, 4–5 µm in diameter. Chemistry: medulla K+faint yellow, C-, PD-. Secondary
metabolites detected by TLC: none. Molecular support: A single ITS sequence (isolate DLH6 from
Haughland 2020-98) has 98% percent identity with Endocarpon unifoliatium GenBank Accession
KX538760 (China, eight positions different, 487 bp overlap, Fig. 3). Given most Endocarpon species have
not been sequenced (NCBI Taxonomy, queried 5 November 2021), we prefer to acknowledge the
uncertainty with “affinity” until further sequences are obtained.
Specimen examined. – CANADA. ALBERTA: Edmonton, Spruce Avenue neighborhood,
53.563562, -113.498138, 2020, weathered and mossy indoor-outdoor carpet, Haughland 2020-98 (hb.
Haughland).
Melanelixia albertana (Ahti) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch
(≡ Melanelia albertana (Ahti) Essl.) FIGURE 37 A.
River valley and ravine system and parkland epiphyte. An occasional boreal species found in
mature mixedwood to deciduous stands. Characterized by an appressed olive-brown to red-brown thallus
with numerous marginal, labriform, and typically downward-facing soralia. The surface often has a shiny,
almost greasy lustre. The lobes are somewhat rounded, and pseudocyphellae absent. One of two fairly
common Melanelixia species in the province, this species can be distinguished from M. subaurifera by the
latter’s laminal soredia intermixed with tiny isidia. Chemistry: medulla C+ red, KC+ red, all other spot tests
negative, lecanoric acid (Esslinger 1977). Molecular support: no sequences generated, low for the species
globally. Leavitt et al. (2016) found poor support for a M. albertana clade composed of sequences from
North America, Russia, China, and India. Regardless of future revisions, our material should retain this
epithet as the type was collected from near Edmonton (Esslinger 1977).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Urban
Monitoring Site 128, 53.614601, -113.439527, 2019, on deciduous tree, A. Hood (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/5454/details); Edmonton, between Stony
Plain Rd. and 100 Ave. at 148 St., 1976, on wooden footbridge, D.C. Lindsay s.n. (PMAE-B77.24.58);
Edmonton, Terwillegar Park, 1977, on Populus bark, D.C. Lindsay s.n. (PMAE-B77.24.83); Edmonton,
Emily Murphy Park, 1976, on Betula bark, D.C. Lindsay s.n. (PMAE-B77.24.90).
Melanelixia subaurifera (Nyl.) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch
(≡ Melanelia subaurifera (Nyl.) Essl.) FIGURE 37 B.
River valley and ravine system epiphyte. A common boreal species with an olive to reddish brown
epruinose thallus. Lobes rounded, flat, usually with both short cylindrical, unbranched isidia mixed with
soredia in laminal and marginal soralia that result in yellow patches where abraded. Pseudocyphellae absent
or inconspicuous; lacking hairs on the lobe tips. Commonly misidentified as Melanelixia subargentifera
Nyl. in Alberta, but that species is rare or absent (ABMI 2020). Chemistry: medulla C+ red, KC+ red, all
other spot tests negative. Secondary metabolites detected by TLC: lecanoric acid. Molecular support: no
sequences generated, inferred to be strong from Leavitt et al. (2016), however, given their description of
cryptic sister species to M. subaurifera, future work should confirm this.
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Figure 37. Brown Parmeliaceae of Edmonton. A, B & D are images of collections from outside of
Edmonton. A, Melanelixia albertana ca. 45 km NE of Beaver Lake, ABMI Site 794, 54.97, -111.34, 2015,
on Populus, C. Copp s.n. [ABMI Lichen # 676192], photo: Dominik Royko. B, Melanelixia subaurifera,
Chickakoo Natural Area west of Edmonton, on Betula, Haughland unvouchered observation. C,
Melanohalea exasperatula, Haughland 2020-24. D, Melanohalea subolivacea, Haughland 2021-36.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Patricia Ravine, 53.503216, -113.592645, 2020, on Prunus stems, D. Haughland & A. Hood (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/14353/details); Edmonton, MacKenzie
Ravine, 1976, on Betula bark, D.C. Lindsay s.n. (PMAE-B77.24.167); Edmonton, approx. 180 St. and 99
Ave., 1976, on twigs and dead branches, D.C. Lindsay s.n. (PMAE-B77.24.70).
Melanohalea exasperatula (Nyl.) O. Blanco, A. Crespo, Divakar, Essl., D. Hawksw. & Lumbsch
(≡ Melanelia exasperatula (Nyl.) Essl.) FIGURE 37 C.
River valley and occasionally tableland epiphyte. Thallus closely appressed, olive-brown and
shiny, but becoming rather rough due to abraded isidia. Lacking pseudocyphellae or soredia. The isidia are
hollow and constricted at the base, initially globular/spherical and becoming flattened to lobulate. There are
several isidiate olive-brown lichens in Alberta that can be difficult to distinguish; the isidia at various
developmental stages are the best diagnostic trait (for drawings see, Esslinger 1977). Chemistry: all spot
tests negative. Secondary metabolites detected by TLC: none. Molecular support: strong at genus- and
species-level even with 105 sequences and circumboreal sampling (Leavitt et al. 2013).
Specimens examined. – CANADA. ALBERTA: Sherwood Park, Air Quality Monitoring Station,
53.532016, -113.321511, 2019, on trunk of Fraxinus pennsylvanica, D. Thauvette & J. Birch s.n. [UoA-
CC-105] (hb. Haughland); Edmonton, Hawrelak trail off-leash park, 53.520696, -113.541533, 2020, on
110
Betula bark, D. Haughland 2020-18 (hb. Haughland); Edmonton, Hawrelak trail off-leash park, 53.519809,
-113.540503, 2020, on Picea glauca twigs, D. Haughland 2020-24 (hb. Haughland); Edmonton,
MacKenzie Ravine, 1976, on Betula, D.C. Lindsay s.n. (PMAE-B77.24.160).
Phaeophyscia kairamoi (Vain.) Moberg FIGURE 38 A-B.
River valley and parkland epiphyte. Like Physcia aff. dimidiata, this species is described in the
southern part of its range as commonly saxicolous (Esslinger 2004). In Alberta it is a common forest
epiphyte, often co-occurring with cyanolichens in mature deciduous or mixedwood stands. Characterized
by irregular thalli that form patches up to 10 cm in diameter, with marginal granular soredia to soredio-
isidia that develop hyaline “hairs,” giving thalli a shaggy look. The upper cortex varies from dark grey to
brown, the lower cortex is black and typically bristling with abundant, simple, white-tipped rhizines. It is
difficult to differentiate from Phaeophyscia hirsuta (Mereschk.) Essl., however P. kairamoi always has
cortical hairs present on the granular soredia, rarely on the lobe ends, whereas P. hirsuta often has cortical
hairs on the lobe ends, but rarely on the soredia (Esslinger 2016a). Molecular support: work is required,
samples submitted to T. Esslinger and S. Leavitt (unpublished data).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 47, 53.505866, -113.553095, 2019, on trunk of Populus balsamifera, D. Haughland s.n. [UoA-CC-86]
(hb. Haughland); Edmonton, Urban Monitoring Site 163E, 53.608013, -113.590864, 2019, on trunk of
Populus balsamifera, D. Haughland & L. Hjartarson s.n. [UoA-CC-84] (hb. Haughland); Edmonton,
Patricia Ravine, 53.504611, -113.593583, 2020, on trunk of live >75 cm DBH Populus balsamifera along
trail, D. Haughland 2020-52 & A. Hood (hb. Haughland); Edmonton, Patricia Ravine, 53.504611, -
113.593583, 2020, on upper branches of downed Populus balsamifera, D. Haughland 2020-56 & A. Hood
(hb. Haughland); Edmonton, MacKenzie Ravine, 1976, on bark of Populus, D.C. Lindsay s.n. (PMAE-as
minor component in B77.24.39).
Phaeophyscia nigricans (Flörke) Moberg FIGURE 38 C-D.
Apparently an uncommon parkland, tableland and river valley epiphyte, but easily overlooked due
to its small size and bark-like color. Characterized by very small lobes <0.5 mm wide on average, forming
thalli up to 1 cm in diameter. Thalli often appear almost sub-fruticose, loosely attached, grey-brown to
brown, blastidiate to soredio-isidiate along margins, rarely developing labriform and continuous marginal
soralia. Distinguished from small, immature Phaeophyscia orbicularis and P. pusilloides (Zahlbr.) Essl. by
size, the white lower cortex and the tiny, hyaline hairs on the lobe tips. Apothecia not observed. Chemistry:
all spot tests negative, no secondary metabolites detected (Esslinger 2004). Molecular support: limited
species-level support (Lohtander et al. 2000), two ITS sequences in GenBank, work is required, sample
submitted to T. Esslinger and S. Leavitt (unpublished data).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 62, 53.520926, -113.432139, 2019, on trunk of Ulmus americana, D. Royko & D. Fielder s.n. [UoA-
CC-15] (hb. Haughland); Edmonton, Urban Monitoring Site 69, 53.534155, 113.525612, 2019, on trunk of
Populus balsamifera, D. Haughland & S. Toni s.n. [UoA-CC-78] (hb. Haughland); Edmonton, Urban
Monitoring Site 86, 53.561777, -113.547997, 2019, on trunk of Ulmus americana, D. Haughland & A.
Hood s.n. [UoA-CC-124] (hb. Haughland); Edmonton, Urban Monitoring Site 160E, 53.583982, -
113.372408, 2019, on trunk of Fraxinus pennsylvanica, S. Toni & M. Cao s.n. [UoA-CC-80] (hb.
Haughland); Edmonton, Urban Monitoring Site 57X, 53.521192, -113.548532, 2019, on trunk of Populus
balsamifera, D. Haughland & A. Hood s.n. [UoA-CC-67] (hb. Haughland); Sherwood Park, Air Quality
Monitoring Station, 53.532016, -113.321511, 2019, on trunk of Fraxinus pennsylvanica, D. Thauvette & J.
Birch s.n. [UoA-CC-105] (hb. Haughland); Edmonton, Urban Monitoring Site 68, 53.530298, -113.554039,
2019, on trunk of Ulmus, D. Haughland s.n. [UoA-CC-17] (hb. Haughland).
Phaeophyscia orbicularis (Necker) Moberg FIGURES 35 C, 38 E-F.
Extremely common tableland, parkland, and river valley epiphyte. A highly variable lichen in
Edmonton and surrounding Parkland and Boreal forests. Characterized by its orbicular thallus (to 3 cm
diameter) and flat, dull lobes, it can form large, confluent patches. The upper cortex is grey-brown to dark
brown, with a black lower cortex, at least centrally. The rhizines tend to be shorter and sparser than in other
111
Figure 38. Phaeophyscia of Edmonton. A-B, P. kairamoi, Haughland 2020-56. A, Thallus. B, Lobe tips
with soredia bearing hyaline hairs. C-D, P. nigricans. C, Thallus, UoA-CC-124. D, Lobe tips with minute
hyaline hairs, UoA-CC-82. E-F, P. orbicularis. E, Thallus, Haughland s.n., photo: Joseph D. Birch. F,
Smooth lobe tips with rhizines visible, UoA-CC-82.
112
Phaeophyscia species. The soralia are greenish, laminal, often circular, but can also be capitate, eroded,
and/or marginal. Often parasitized and then with lumpy, laminal darkened outgrowths. Apothecia
occasional, and then with sparse rhizines on the thalline margin. Shade forms with a whitish distal lower
cortex can be confused with Physciella melanchra (Hue) Essl. but that species has a uniformly pale
prosoplectenchymatous lower cortex (vs. paraplectenchymatous and dark at least centrally in P.
orbicularis). To date we have not confirmed that P. melanchra occurs in our region. Chemistry: all spot
tests negative, no secondary metabolites detected (Esslinger 2004). Molecular support: limited species-level
support (Lohtander et al. 2000), work is required, samples submitted to T. Esslinger and S. Leavitt
(unpublished data).
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring
Site 6, 53.454045, -113.595336, 2019, on trunk of Populus sp., D. Haughland & A. Stordock s.n. [UoA-
CC-142] (hb. Haughland); Edmonton, Urban Monitoring Site 40, 53.493790, -113.504979, 2019, on trunk
of Fraxinus pennsylvanica, D. Haughland & M. Cao s.n. [UoA-CC-8,16] (hb. Haughland); Edmonton,
Urban Monitoring Site 62, 53.520684, -113.432139, 2019, on trunk of Ulmus americana, D. Royko & D.
Fielder s.n. [UoA-CC-157] (hb. Haughland); Edmonton, Urban Monitoring Site 95, 53.570645, -
113.588770, 2019, on trunk of Ulmus americana, M. Lewis & M. Villeneuve s.n. [UoA-CC-166] (hb.
Haughland); Edmonton, Urban Monitoring Site 107E, 53.548249, -113.617818, 2019, on trunk of
Elaeagnus angustifolia, D. Haughland s.n. [UoA-CC-9] (hb. Haughland); Edmonton, Urban Monitoring
Site 148E, 53.587522, -113.640605, 2019, on trunk of Populus balsamifera, D. Haughland & L.
Hjartarson s.n. [UoA-CC-48] (hb. Haughland); Edmonton, between Provincial Museum parking lot and
Wellington Crescent, 1975, on N sides of trunks of deciduous trees, D.C. Lindsay s.n. (PMAE-B77.24.47);
Edmonton, McKinnon Ravine, 1976, on tree bark, D.C. Lindsay s.n. (PMAE-B77.24.50); Edmonton,
Terwillegar Park, 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.80); Edmonton, Whitemud Park, 1976,
on wood, D.C. Lindsay s.n. (PMAE-B77.24.126); Edmonton, along Saskatchewan Drive, near the
Biological Sciences Building of the University of Alberta, 1976, on bark (cf. Salix), D.C. Lindsay s.n.
(PMAE-B77.24.175).
Physconia detersa (Nyl.) Poelt FIGURE 39 A.
Occasional river valley epiphyte. The squarrose rhizines and pruinose lobes are key in separating
Physconia from similar genera. Characterized by dull, pruinose grey-brown to dark brown lobes, thalli may
grow up to 6 cm in diameter. The lower cortex is well-developed and black with abundant squarrose
rhizines. The upper cortex is scleroplectenchymatous. The medulla is white, and the soralia marginal.
Physconia enteroxantha is similar in appearance, differing in its pale-yellow medulla that reacts K+ yellow,
KC+ dark yellow. Chemistry: all spot tests negative, no secondary metabolites detected normally, variolaric
acid may be an accessory in soralia (Esslinger 2002). Molecular support: a single ITS sequence (isolate
DLH17 from Haughland 2020-51) is 100% identical to P. detersa GenBank Accession EF582760 (Finland,
486 bp overlap) and 99% identical to KT695314 (Canada, Ontario, two different positions, 498 bp overlap).
Monophyletic with low support in Esslinger et al. (2017). In our analyses, a GenBank sequence of
Physconia jacutica nests within an otherwise well-supported monophyletic clade of P. detersa (Fig. 8).
Specimens examined. – CANADA. ALBERTA: Edmonton, Patricia Ravine, 53.504611, -
113.593583, 2020, on trunk of live >75 cm DBH Populus balsamifera along trail, D. Haughland 2020-51
& A. Hood (hb. Haughland); Edmonton, Patricia Ravine, 53.504611, -113.593583, 2020, on upper branches
of downed Populus balsamifera, D. Haughland 2020-57 & A. Hood (hb. Haughland); Edmonton,
MacKenzie Ravine, 1976, on Betula bark, D. C. Lindsay s.n. (PMAE-B77.24.155); Edmonton, MacKenzie
Ravine boardwalk, 53.528875, -113.558827, 2021, on decayed Picea glauca log, D. Haughland 2021-10
(hb. Haughland); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315, 2021, on Betula papyrifera, D.
Haughland 2021-22A (hb. Haughland).
Physconia enteroxantha (Nyl.) Poelt FIGURE 39 B.
Occasional river valley epiphyte. Thalli to 5 cm diameter, closely attached, green-brown to dark
brown, often with white pruina. The soralia are marginal, rarely labriform, occasionally becoming laminal,
and the soredia can become lobulate centrally, verging on Physconia grumosa Kashiw. & Poelt. The lower
cortex is brown to black with abundant squarrose rhizines. Medulla and soralia typically are yellowish.
Distinguishing this species from P. grumosa and P. detersa is difficult when secondary metabolite
113
Figure 39. Physconia of Edmonton. A, P. detersa, Haughland 2020-51. B, P. enteroxantha, wet,
Haughland 2020-29. The following two images are of collections are from outside of Edmonton. C, P.
labrata thallus, dry, William Switzer Provincial Park, 2015, on Picea glauca, Haughland 2015-1a. D, P.
perisidiosa thallus, dry, ca. 40 km SW of Fort Chipewyan, ABMI Site 208, 58.49, -111.69, 2010, on
Populus, T. Grainger s.n. [ABMI Lichen # 247465].
concentrations are low, but these species have a scleroplectenchymatous upper cortex while P.
enteroxantha is paraplectenchymatous (Esslinger 2016c). This is best seen in longitudinal sections about 2–
3 mm from the lobe tips. In addition, P. grumosa has more granular, scattered soredia, forming branched
isidioid soredia and lobules centrally. Chemistry: medulla and soralia K+ yellow, KC+ dark yellow, all
other spot tests negative. Secondary metabolites detected by TLC: none, despite analyzing 11 specimens
with positive K spot tests. The amount of secalonic acid A in the medulla and soralia can vary greatly
(Esslinger 2002). Molecular support: A single ITS sequence (isolate DLH32 from Haughland 2020-29) is
99% identical to Ph. enteroxantha GenBank Accessions LS483215 (Spain, one position different, 697 bp
overlap) and MK811936 (Norway, two positions different, 488 bp overlap). Monophyletic in Esslinger et
al. (2017) and our analyses (Fig. 8).
Specimens examined. – CANADA. ALBERTA: Edmonton, South Air Quality Monitoring
Station, 53.501946, -113.5249, 2019, on trunk of Fraxinus sp., D. Thauvette & J. Birch s.n. [UoA-CC-132]
(hb. Haughland); Edmonton, Hawrelak trail off-leash park, 53.520696, -113.541533, 2020, on Betula bark,
D. Haughland 2020-23 (hb. Haughland); Edmonton, Kinnaird Ravine, 53.558953, -113.459253, 2020, on
Picea glauca snag, 2020, D. Haughland 2020-29 & P. Williams (hb. Haughland); Edmonton, MacKenzie
Ravine, 1976, on bark of Populus, D.C. Lindsay s.n. (PMAE-B77.24.39); Edmonton, Terwillegar
Footbridge, 53.483201, -113.600184, 2021, on Betula papyrifera, D. Haughland 2021-12 (hb. Haughland).
114
Physconia labrata Essl., McCune & Haughland FIGURE 39 C.
Rare river valley epiphyte. A recently described species (Esslinger et al. 2017), formally
considered part of the Physconia perisidiosa species concept. Both species are pruinose, form well-
developed rosette-like thalli to sparse, poorly developed shingle-like thalli, and develop labriform, terminal
soralia. Physconia labrata differs in its well-developed dark lower cortex (vs. poorly-developed lower
cortex, pale with sparse, dark hyphae in P. perisidiosa) and paraplectenchymatous upper cortex (vs.
scleroplectenchymatous in P. perisidiosa). Relatively abundant in the Boreal and Foothills Natural Region
(ABMI 2020, Esslinger et al. 2017). Chemistry: all spot tests negative. Secondary metabolites detected by
TLC: none. Molecular support: high species-level support, type from region of Hinton, Alberta,
reciprocally monophyletic in Esslinger et al. (2017; Fig. 8 herein), no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Patricia Ravine, 53.504611, -
113.593583, 2020, on trunk of live >75 cm DBH Populus balsamifera along trail, D. Haughland 2020-53
& A. Hood (hb. Haughland).
Physconia perisidiosa (Erichsen) Moberg FIGURE 39 D.
Rare river valley epiphyte. Like Physconia labrata but with a scleroplectenchymatous upper
cortex and a poorly developed fibrous lower cortex (Esslinger et al. 2017). Chemistry: all spot tests
negative. Secondary metabolites detected by TLC: none. Molecular support: forming a well-supported but
unresolved clade ith P. venusta (Ach.) Nyl. (Esslinger et al. 2017; Fig. 8 herein), no new sequences
generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, ravine between Stony Plain Rd. and
100 Ave. at 148 St., 1976, on tree bark, D.C. Lindsay s.n. (PMAE-B77.24.52).
GROUP 11. SMALL CYANOLICHENS
Four species. Key literature: Goward et al. 1994; Jørgenson & Nash 2004; Otáloro et al. 2008,
2014; Schultz & Büdel 2002. We did not document chemistry as it is not typically useful for identifying
members of this group.
1a. Tiny gelatinous granules growing on concrete, hardly visible without >10x magnification; spores
simple ................................................................................................................................... Lichinaceae sp. 1
1b. Small foliose to semi-fruticose thalli visible with the naked eye (but often camouflaged and difficult to
detect) growing on soil; spores transversely septate to muriform ................................................................... 2
2a. Appearing dwarf-fruticose but with foliose lobes at least proximately; apothecia abundant; lobes with
cellular cortices visible in cross-section ..................................................................... Scytinium tenuissimum
2b. Lobes clearly foliose; apothecia present or absent; lobes lacking cortices in cross-section ...................... 3
3a. Lobes margins swelling disproportionately when moist; spores muriform ..................... Enchylium tenax
3b. Lobes swelling evenly when moistened; spores with up to 3 transverse septa..... Blennothallia crispa s.l.
Blennothallia crispa (Hudson) Otálora, P. M. Jørg. & Wedin s.l.
(≡ Collema crispum (Hudson) Weber ex F. H. Wigg) FIGURE 40 A-C.
River valley terricole. A relatively common lichen in southern Alberta (ABMI 2020). The
Edmonton specimen is a rare example of fertile material. Edmonton material: lobes overlapping, isidia-like,
semi-fimbriate, appressed, margins not swollen disproportionately when wet. Apothecia constricted at the
base, spores with transverse septa, up to 4-celled, 25–35 × 10–12 µm, 8 per ascus. Apothecium with
lobulate proper exciple, lobes ecorticate with heteromerous interior of elongated hyphae oriented at right
angles, with small-celled Nostoc in short chains or clusters. Molecular support: uncertain. A single ITS
sequence (isolate DLH31 from Haughland 2020-60) did not have query coverage >75% with any GenBank
accessioned sequences, and of those with coverage 50–75%, no sequence exceeded 91% percent identity.
115
We did not find any accessioned ITS sequences for this species through NCBI Taxonomy Browser;
additional loci and comparative reference sequences are required for phylogenetic analyses.
Specimen examined. – CANADA. ALBERTA: Edmonton, Hawrelak trail off-leash park,
53.517625, -113.5402, 2020, on exposed mineral soil and moss, D. Haughland 2020-60 & Kyla Tichkowski
(hb. Haughland).
Enchylium tenax (Sw.) Gray
(≡ Collema tenax (Sw.) Ach.) FIGURE 40 D-E.
River valley terricole. Edmonton material: small black appressed lobes, immature lobes appearing
globular. Apothecia forming laminally, spores hyaline, muriform, 25–30 × 11–18 µm. Nostoc in thallus
forming long chains. Edmonton material corresponds morphologically to what was previously called
Collema tenax var. crustaceum (Kremp.) Degel. (Goward et al. 1994). Molecular support: no new
sequences generated; species paraphyletic with E. polycarpon (Hoffm.) Otálora, P. M. Jørg. & Wedin in
recent phylogeny (Otálora et al. 2014).
Specimen examined. – CANADA. ALBERTA: Edmonton, Emily Murphy Park, near LRT bridge
and Kinsmen Sports Centre, 53.527438, -113.514725, 2020, on moist N-facing mineral soil slope toe, along
trail, D. Haughland 2020-39 & P. Williams (hb. Haughland).
***Lichinaceae sp. 1 FIGURE 41.
Growing on old, exposed, rough concrete in the inner city, with Caloplaca feracissima and
Caloplaca tominii, these tiny thalli may belong to the genus Lichinella or Gonohymenia (M. Schultz, pers.
comm.); more work is needed. Thalli consist of aggregates of cyanobacteria that form small granules with a
polygonal surface in squash, with sparse hyphae visible in cross-section. The smallest of these granules are
35–25 × 25 µm, and some thalli consist solely of discrete tiny granules while others form larger
aggregations. The larger aggregations regularly contain fruiting bodies that are all but invisible even under
40× magnification and appear to be embedded in the thallus with a layer of algae overlying the fruiting
bodies. Upon squash or cross-section, asci can be found, apparently forming thallinocarps (Schultz & Büdel
2002). At maturity the asci are clavate, 8-spored, 18 × 35 µm. No paraphyses could be identified. The walls
of mature asci stain K/I+ blue, and immature asci appear to show either a Lecanora-type or Fuscidea-type
internal structure, with an amyloid tip. The spores are simple, hyaline, ovoid to bean-shaped, in rare
instances slightly constricted in the middle or curved, 10–12 × 5–6 µm, and they stain yellow in K/I. The
conidia are bacilliform, 2–4 × 1 µm, and are formed from unbranched conidiophores in globose, embedded
pycnidia. The tissues showed no reaction to K or C alone as tested by drawing the chemicals under a cover
slip while observing with a compound microscope. The photobiont could not be determined with certainty
and appeared to be a mix of trebouxioid, Chroococcus-type and single-celled cyanobacteria, with some in
clusters of 2–5 cells. Molecular support: none, a single specimen failed to amplify.
Specimen examined. – CANADA. ALBERTA: Edmonton, Spruce Avenue Neighborhood,
53.563562, -113.498138, 2020, on old concrete, D. Haughland 2020-95C (hb. Haughland).
Scytinium tenuissimum (Dickson) Otálora, P. M. Jørg. & Wedin
(≡ Leptogium tenuissimum (Dickson) Körber) FIGURE 40 F-H.
Apparently rare river valley terricole. Growing on moist, trailside mineral soil and moss alongside
Peltigera and Cladonia. Thallus composed mostly of fimbriate, coralloid isidia developing from foliose
lobes that were difficult to find and may have been senescing. The dominance of the isidia was suggestive
of Scytinium teretiusculum (Wallr.) Otálora, P. M. Jørg. & Wedin; however, the abundant apothecia and
apothecial morphometrics support the placement of this collection in S. tenuissimum. Apothecia abundant,
thalline margins becoming isidiate, disk reddish brown, concave to almost plane, to 1 mm in diameter.
Hymenium 170 µm thick, spores hyaline, muriform, 29–35 × 12–15 µm. Molecular support: no new
sequences generaqted. In Otálora et al. (2014) two sequences form a well-supported branch nested within a
clade containing S. subtile (Schrad.) Otálora, P.M. Jørg. & Wedin and S. palmatum (Hudson) Gray. More
work is needed.
116
Figure 40. Terricolous cyanolichens of Edmonton. A-C, Blennothallia crispa s.l., Haughland 2020-60. A,
Thallus in the field, wet, inset showing lobes and apothecium under magnification. B, Lobe cross-section
showing largely heteromerous interior of elongated hyphae oriented at right angles, lack of cortices and
short chains of Nostoc. C, Transversely-septate spores within asci, same scale as B. D-E, Enchylium tenax,
Haughland 2020-39. D, Thallus in the field, wet. E, Muriform spores within asci. F-H, Scytinium
tenuissimum, Haughland 2021-14 https://naturelynx.ca/sightings/17278/details. F, Thallus in the field. G,
Spores in ascus. H, Isidiate lobe tips.
117
Figure 41. Concrete-dwelling Lichinaceae sp. 1 from Edmonton, Haughland 2020-95C. A, Macroscopic
view of wet thallus, ranging from individual granules to aggregates. B, Pycnophores embedded in thallus
and bacilliform conidia. C, Asci and spores in water-mount D, Individual algal granules in water. E-F, Asci
after treatment with K and Lugol’s Iodine.
118
Specimen examined. – CANADA. ALBERTA: Edmonton, Fort Edmonton Park region,
53.500919, -113.576036, 2021, on trailside mineral soil and moss, D. Haughland 2021-14 & S. Toni (hb.
Haughland).
GROUP 12. PELTIGERA
Eleven species. An ongoing collaboration with F. Lutzoni, J. Miadlikowska, T. Goward, I.
Medeiros and C. Pardo-De la Hoz has confirmed that Alberta is home to many morphologically similar,
undescribed species of Peltigera (sensu Miadlikowska et al. 2003, 2018; Pardo-De la Hoz et al. 2018).
With few Edmonton specimens sequenced, here we use currently accepted morphological species concepts
as described in Goffinet and Hastings (1994) and Goward et al. (1995) as well as molecular species where
available from Pardo-De la Hoz et al. 2018 (section Chloropeltigera) and Magain et al. 2018 (section
Peltigera). Nature-recording app records exist for P. ponojensis Gyelnik, but images and collected material
were insufficient to confirm this species for Edmonton. Because we have not used chemistry to distinguish
species to date, we have not included references to secondary metabolites documented elsewhere.
1a. Tripartite “freckled” lobes, primary photobiont a green alga giving lobes a bright green color when
hydrated, secondary photobiont the cyanobacterium Nostoc confined to wart-like cephalodia over the upper
surface; a diverse group of which only a single species is confirmed in Edmonton to date ..............................
...................................................................................................................................... Peltigera leucophlebia
1b. Bipartite, lacking “freckle”-like cephalodia, primary photobiont the cyanobacterium Nostoc; lobes grey
to brown to almost black ................................................................................................................................. 2
2a. Soredia present, in laminal circular soralia ................................................................................................ 3
2b. Soredia absent ........................................................................................................................................... 4
3a. Lobes deeply concave, mostly growing singly; rhizines simple, sparse ....................... Peltigera didactyla
3b. Lobes often polyphyllous; rhizines flocculent, abundant ............................................ Peltigera extenuata
4a. Laminal isidia present, isidia mostly granular, cylindrical or corraloid; lobules typically absent ...............
.......................................................................................................................................... Peltigera evansiana
4b. Laminal isidia absent; lobules may be present along edges of cracks, scars, or along lobe edges ............ 5
5a. Upper cortex with spider web-like arachnoid tomentum ........................................................................... 6
5b. Upper cortex smooth, scabrid (with a scabby crust of crystals, dead cells and tomentum) or pruinose,
but not tomentose .......................................................................................................................................... 10
6a. Lobe tips mostly upturned and concave .................................................................................................... 7
6b. Lobe tips mostly downturned or ruffled, like an Elizabethian collar ........................................................ 8
7a. Thalli asymmetrical, like small ascending hands; finger-like lobes bearing apothecia that ≤6 mm in
length; soralial scars often present in center of thallus; upper surface with patchy, thin tomentum, typically
lacking pruina; rhizines simple, discrete .................................................... Peltigera didactyla, fertile morph
7b. Thalli often symmetrical, radiating lobes appressed throughout; when fertile, finger-like lobes bear
apothecia ≤12 mm in length; lacking soralial scars; upper surface typically tomentose throughout, with
pruina crusting central portions; rhizines dense, in hedgerows, becoming confluent centrally .........................
........................................................................................................................................... Peltigera rufescens
8b. Lobes relatively large and wedge-shaped, with consistently down-turned edges; upper surface
tomentose throughout; rhizines mostly squarrose-branched, often becoming confluent ........ Peltigera canina
8b. Lobes variable in size, edges ruffled; upper tomentum often patchy throughout or restricted to the outer
lobe surface; rhizines mostly discrete .............................................................................................................. 9
119
9a. Upper surface often with wiggly pale lines from invertebrate grazing; lobe edges and surface stress
cracks forming least some lobules (check centrally); veins raised, appearing overlapping near edges, often
pinkish to cinnamon brown in color; rhizines mostly simple and smooth; a hypervariable species
morphologically, overlapping with traditional concepts of P. membranacea and P. ponojensis ......................
......................................................................................................................................... Peltigera praetextata
9b. Upper surface typically ungrazed; lobules typically absent; veins raised but not overlapping, becoming
chocolate brown to dark brown to black centrally; rhizines simple but developing some squarrose
branching; difficult to confidently discriminate with morphology alone from related section Peltigera
species ................................................................................................................................ Peltigera islandica
10a. Upper surface grey, typically somewhat shiny, and often with rows of slight depressions or “dimples”
across the lobes; lower surface with rhizines aligned in concentric rows and indistinct, broad, flat veins;
when fertile, apothecia held flat and more or less parallel to lobe surface .................................................... 11
10b. Upper surface grey to deep green; lower surface with rhizines not aligned and typically with distinct
veins; when fertile, apothecia forming tight “hot dog bun” rolls that are held upright ................................. 12
11a. Upper surface typically with numerous stress cracks, developing abundant lobules and schizidia; veins
dark brown to almost black, indistinct or apparently absent, interstices between veins shallow and rounded;
common ........................................................................................................................... Peltigera elisabethae
11b. Upper surface lacking stress cracks and lobules; veins medium brown, broad to occasionally
indistinct; interstices more or less numerous, often elongate; rare in Alberta ............... Peltigera horizontalis
12a. Upper surface steel grey to grey-blue when hydrated, often lightly pruinose, lacking lobules; apothecia
black; veins abruptly darkening toward center; common ...................................................... Peltigera neckeri
12b. Upper surface deep green when hydrated, lacking pruina but often with lobules; apothecia reddish-
brown, forming at the tips of long, ascending, deeply dissected lobes; veins reddish-brown to dark brown;
rare ........................................................................................................ Peltigera polydactylon subsp. udeghe
Peltigera canina (L.) Willd. FIGURE 42 A-B.
River valley and ravine system and rare tableland terricole. A taxonomically challenging species
with many undescribed species currently included within—applied in the broad sense here to thalli that
have large, downturned lobes, extensive tomentum on the upper surface, and squarrose to flocculent
rhizines that become matted below. Lacking soredia, isidia or lobules. This is the only species of Peltigera
found in a lawn niche outside of the river valley parks. Molecular support: two ITS sequences (isolate
DLH24 from Haughland 2019-120, isolate DLH30 from Haughland 2020-7), are >99% similar to
specimens corresponding to Peltigera canina 2, and they contain the canina 2 hypervariable region from
Magain et al. (2018; Fig. 7 herein).
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.502638, -
113.602538, 2020, on debris and downed wood, D. Haughland 2020-7 & C. Shier (hb. Haughland);
Edmonton, Alberta Avenue neighborhood, 53.564178, -113.490616, 2019, terricolous in mowed grass
along boulevard, D. Haughland 2019-120 & P. Williams (hb. Haughland); Edmonton, near Northland
sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on soil, D.C. Lindsay s.n. (PMAE accession
B77.24.102).
Peltigera didactyla (With.) J. R. Laundon FIGURE 43 A-B.
Apparently rare river valley terricole. Elsewhere in Alberta this is a common early successional
species (ABMI 2020). Characterized by small, deeply concave lobes with round, laminal soralia, and
relatively sparse, long, simple rhizines below. This species intergrades with Peltigera extenuata and the
under-detected and often misidentified P. castanea Goward, Goffinet & Miądl. in many regions of Alberta
(ABMI 2020). Molecular support: one sample failed to amplify cleanly enough to sequence. The difficulty
diagnosing these morphologically similar species means this species is a priority for future sequencing.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
53.502638, -113.602538, 2020, terricolous, D. Haughland 2020-9 & C. Shier (hb. Haughland); Edmonton,
west of Whitemud Park, along Grandview Stairs, 53.502565, -113.553934, 2020, on moss, L. Hjartarson
(unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/12369/details); Edmonton,
MacKenzie Ravine, 1976, on soil, D.C. Lindsay s.n. (PMAE accession B77.24.43).
120
Figure 42. Tomentose Peltigera species of Edmonton, plate 1 of 2. Collections imaged in C-D are from
outside of Edmonton. A-B, P. canina. A, Habit, Haughland unvouchered observation. B, Lower surface
showing raised veins and flocculent rhizines, Haughland 2020-7. C-D, P. islandica, ca. 10 km SE of
Cadomin, ABMI Site 1232, 52.97, -117.22, 2011, on downed wood, M. Martel s.n. [ABMI Lichen #
292500], identification verified using ITS. C, Upper surface. D, Lower surface with dark veins and discrete
rhizines. E-F, P. rufescens. E, Habit, Haughland 2020-38. F, Lower surface with confluent rhizines,
Haughland 2021-17.
121
Figure 43. Sorediate and isidiate Peltigera species of Edmonton. A-B, P. didactyla, Laura Hjartarson,
https://naturelynx.ca/sightings/12369/details. A, Habit and soralia. B, Fertile thallus with soralia. C-D, P.
evansiana, Haughland 2020-36. C, Habit. D, Isidia on upper surface. E-F, P. extenuata, Haughland 2020-
21. E, Habit and soralia. F, Rhizines on lower surface.
122
Peltigera elisabethae Gyelnik FIGURE 44 A-B.
River valley and ravine system xylicole/terricole. One of Edmonton’s most common lichens,
forming extensive colonies (>1 m2) over soil and downed wood. Characterized by large, often shiny grey
lobes that lack tomentum, marginal lobules and stress cracks that result in schizidia, flat red-brown
apothecia, rhizines aligned in rows, and round interstices on a largely veinless lower surface. Molecular
support: one ITS sequence (isolate DLH25 from Haughland 2020-12) is 95% identical to P. elisabethae
GenBank Accession MK517830; Fig. 7 herein).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf
Willow Creek, 53.501775, -113.6017, 2020, on mossy slope, D. Haughland 2020-12 & C. Shier (hb.
Haughland); Edmonton, Larch Sanctuary, 53.452705, -113.547888, 2020, terricolous, D. Haughland
(unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/10686/details); Edmonton,
Fort Edmonton Park, 53.499113, -113.58172, 2020, on moss, cwf_michelleh (unvouchered observation:
iNaturalist record https://www.inaturalist.org/observations/58992454); Edmonton, MacKenzie Ravine,
1976, on soil, D.C. Lindsay s.n. (PMAE-B77.24.28); Edmonton, MacKenzie Ravine, 1976, on damp soil,
D.C. Lindsay s.n. (PMAE-B77.24.171); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315, 2021,
on moss, D. Haughland 2021-13 & S. Toni (hb. Haughland).
Peltigera evansiana Gyelnik FIGURE 43 C-D.
River valley and ravine system xylicole/terricole. Frequent in mature mixedwoods, this species is
unmistakable due to its combination of large, down-turned, tomentose lobes and abundant laminal, granular
isidia. The lower surface resembles Peltigera praetextata, with narrow, raised beige veins and simple
rhizines. Molecular support: no new sequences generated; species-level support high (Magain et al. 2018).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Emily
Murphy Park, near LRT bridge and Kinsmen Sports Centre, 53.52742, -113.514617, 2020, on moist north-
facing mineral soil slope toe, along trail, D. Haughland 2020-36 & P. Williams (hb. Haughland,
NatureLynx record https://naturelynx.ca/sightings/13878/details); Edmonton, Wolf Willow Creek Ravine,
53.5022, -113.60123, 2020, on moss/soil, D. Haughland & C. Shier (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/14355/details); Edmonton, MacKenzie Ravine, 1976, on soil, D.C.
Lindsay s.n. (PMAE-B77.24.29); Edmonton, south bank of North Saskatchewan River, opposite 114 St. at
Saskatchewan Drive, 1986, on side of trail, J.E. Marsh 1647 (PMAE-B86.141.51).
Peltigera extenuata (Vain.) Lojka FIGURE 43 E-F.
Apparently rare river valley terricole. Elsewhere in Alberta this is common on rotten wood (ABMI
2020). Characterized by small, concave, often-polyphyllous lobes with round, laminal soralia, and
abundant, pale, flocculent rhizines below. This species intergrades with Peltigera didactyla and the under-
detected and often misidentified P. castanea in many regions of Alberta (ABMI 2020). Molecular support:
no new sequences generated, the difficulty diagnosing these morphologically similar species means this
species is a priority for sequencing.
Specimen examined & observation. – CANADA. ALBERTA: Edmonton, River Loop Trail S of
Fort Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland 2021-21 & S. Toni (hb.
Haughland, NatureLynx record https://naturelynx.ca/sightings/17282/details).
Peltigera horizontalis (Hudson) Baumg. FIGURE 44 C-D.
River valley xylicole/terricole. Intergrading with Peltigera elisabethae but far rarer than the latter
in Alberta (ABMI 2020). Characterized by the large, smooth, grey, shiny dimpled upper surface, and
aligned rhizines below. Discriminated from P. elisabethae by its lack of lobules and more numerous,
elongate interstices that may form distinct veins. Historical records for the province have largely been
revisited and redetermined as other species. Molecular support: a tentatively identified extant collection
was sequenced but it grouped with P. elisabethae. The difficulty diagnosing these morphologically similar
species means this species continues to be a priority for sequencing.
Specimen examined. – CANADA. ALBERTA: Edmonton, Whitemud Ravine, 1993, on moss, R.
Hastings s.n. (PMAE accession C93.6.3).
123
Figure 44. Glabrous Peltigera species of Edmonton. Collections imaged in C-D are from outside of
Edmonton. A-B, P. elisabethae. A, Fertile colony with abundant lobules, D. Haughland, unvouchered
observation. B, Lower surface showing poorly-defined veins and aligned rhizines, Haughland 2020-12. C-
D, P. horizontalis, ca. 51 km NE of Sandy Lake, ABMI Site 664, 55.85, -112.60, 2015, on downed wood,
S. Venskaitis s.n. [ABMI Lichen # 677340], identification verified using ITS. C, Upper surface. D, Lower
surface showing elongate interstices and aligned rhizines E-F, P. neckeri. G, Fertile colony with hotdog
bun, black apothecia, Haughland https://naturelynx.ca/sightings/17275/details. H, Lower surface with well-
defined black veins, Haughland 2020-59. G-H, P. polydactylon subsp. udeghe. G, Fertile colony,
Haughland 2020-2. H, Lower surface showing well-defined brown veins, Haughland 2020-50.
124
Figure 45. Peltigera leucophlebia, Edmonton. A, Habit, Haughland 2020-34. B, Lower surface with well-
delimited veins, D. Haughland, https://naturelynx.ca/sightings/17274/details.
*Peltigera islandica T. Goward & S.S. Manoharan-Basil FIGURE 42 C-D.
River valley xylicole/terricole. This species appears to be relatively rare in Alberta in general, and
Edmonton in particular (ABMI 2020). Characterized by broad thin lobes, with thin tomentum sometimes
limited to the lobe tips, distinct veins that darken to chocolate brown or black, and typically simple rhizines
that may become flocculent in part. It intergrades with the more common Peltigera canina and P.
praetextata, as well as the much rarer P. membranacea (Ach.) Nyl. in Alberta; work is ongoing to
determine if high fidelity traits exist to differentiate these species. Molecular support: a single ITS sequence
(isolate DLH21 from Haughland 2020-1) is 100% identical to P. islandica GenBank Accession KJ413244
and has the P. islandica/sp 20 hypervariable region from Magain et al. (2018; Fig. 7 herein).
Specimen examined. – CANADA. ALBERTA: Edmonton, Emily Murphy Park, near LRT bridge
and Kinsmen Sports Centre, 53.52742, -113.514617, 2020, on moist N-facing mineral soil slope toe, along
trail, D. Haughland 2020-37A & P. Williams (hb. Haughland).
Peltigera leucophlebia (Nyl.) Gyelnik FIGURE 45.
River valley terricole. This broad-lobed, emerald-green, ruffled species is Edmonton’s only
confirmed cephalolichen. The cephalodia typically are abundant on the upper surface and are rounded to
bulbous. The lower surface has distinct veins and rhizines. When fertile, there are distinct patches of cortex
on the underside of the marginal, red-brown, saddle-shaped apothecia (vs. a more continuous cortex on the
back of apothecia in the more boreal P. aphthosa (L.) Willd.). Molecular support: a single ITS sequence
(isolate DLH20 from Haughland 2020-34) has 99% percent identity to P. leucophlebia 2 GenBank
Accessions MH734662 and MH734664 from Pardo-De la Hoz et al. (2018; Fig. 7 herein).
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Emily Murphy Park,
near LRT bridge and Kinsmen Sports Centre, 53.52742, -113.514617, 2020, on moist N-facing mineral soil
slope toe, along trail, D. Haughland 2020-34 & P. Williams (hb. Haughland, NatureLynx record https://
naturelynx. ca/sightings/13874/details); Edmonton, Rio Park, 53.502290, -113.592075, 2020,
mathew_specht (unvouchered observation: iNaturalist record https://www.inaturalist.org
/observations/42698504); Edmonton, MacKenzie Ravine, 1979, on soil, D.C. Lindsay s.n. (PMAE-
B77.24.27); Edmonton, Whitemud Park, 1976, D.C. Lindsay s.n. (PMAE accession B77.24.108);
Edmonton, River Loop Trail S of Fort Edmonton, 53.500627, -113.576611, 2021, on moist trail-side soil,
D. Haughland & S. Toni (unvouchered observation: Naturelynx record https://naturelynx.ca/
sightings/17274/details).
125
Peltigera neckeri Hepp ex Müll. Arg. FIGURE 44 E-F.
River valley and ravine system xylicole/terricole. Frequent in Edmonton’s river valley parks, this
shiny, grey, etomentose species is differentiated from the more common Peltigera elisabethae by the
tightly curled black “hot dog bun”-shaped apothecia (vs. flat red-brown apothecia in P. elisabethae),
darkened rhizines that are not aligned (vs. aligned in P. elisabethae), and lack of lobules (vs. often
abundantly lobulate P. elisabethae). While variable, this species typically has dark, clearly differentiated
veins below. Molecular support: a single ITS sequence (isolate DLH23 from Haughland 2020-59) has
100% percent identity to P. neckeri GenBank Accession AF075725 (Fig. 7).
Specimens examined. – CANADA. ALBERTA: Edmonton, Emily Murphy Park, near LRT
bridge and Kinsmen Sports Centre, 53.527, -113.51442, 2020, epixylic on decayed wooden bridge edge, D.
Haughland 2020-31 & P. Williams (hb. Haughland, NatureLynx record
https://naturelynx.ca/sightings/13876/details); Edmonton, Wolf Willow Creek, 53.502811, -113.602225,
2020, on downed wood, D. Haughland 2020-4 & C. Shier (hb. Haughland); Edmonton, Patricia Ravine,
53.503811, -113.593841, 2020, on wood in moist depression, D. Haughland 2020-48 & A. Hood (hb.
Haughland); Edmonton, Patricia Ravine, 53.504141, -113.59432, 2020, on downed wood, Haughland
2020-59 & A. Hood (hb. Haughland); Edmonton, River Loop Trail S of Fort Edmonton, 53.500627, -
113.576611, 2021, on moist trail-side soil, D. Haughland & S. Toni (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/17275/details).
Peltigera polydactylon (Necker) Hoffm. subsp. udeghe Magain, Miadl. & Sérus. FIGURE 44 G-H.
River valley xylicole/terricole. Like Peltigera horizontalis, this species appears to be uncommon
across much of Alberta, and is typically found in mature moist and/or riparian forests. It is characterized by
red-brown, tightly curled, erect apothecia on elongate lobes that typically have marginal lobules. Peltigera
polydactylon subsp. udeghe was reported in Edmonton by Magain et al. (2016, B. Goffinet 487, herb. B.
Goffinet[n.v.]; DNA-N1885). Molecular support: a single ITS sequence (isolate DLH26 from Haughland
2020-2) has 100% percent identity to P. polydactylon subsp. udeghe GenBank Accession KX365430 from
Magain et al. (2016; Fig. 7 herein).
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.5028, -
113.602075, 2020, on moss, D. Haughland 2020-2 & C. Shier (hb. Haughland); Edmonton, Patricia
Ravine, 53.50388, -113.594428, 2020, on downed wood, D. Haughland 2020-50 & A. Hood (hb.
Haughland).
Peltigera praetextata (Flörke ex Sommerf.) Zopf FIGURE 46.
River valley xylicole/terricole. Common across Alberta and in Edmonton, but phenotypically
plastic and therefore difficult to differentiate from undescribed Peltigera canina group species, P.
membranacea, P. ponojensis and P. islandica. Typical morphs have ruffled margins and simple, smooth
rhizines with distinct beige veins below. The tomentum is typically restricted to the distal third of the lobes.
A key feature found with diligent searching in the majority of specimens is lobules that form along cracks
and to a lesser extent along lobe edges, although they should not be viewed as definitive evidence of P.
praetextata—P. islandica and P. wulingensis L.F. Han & S.Y. Guo can also be lobulate in Alberta.
Molecular support: four ITS sequences (isolates DLH22 from Haughland 2020-37B, DLH28 from
Haughland 2020-1 [originally identified as P. membranacea], DLH18 from Haughland 2020-35, and
DLH29 from Haughland 2020-58) have 100% percent identity to published P. praetextata GenBank
accessioned sequences as well as the P. praetextata hypervariable region from Magain et al. (2018; Fig. 7
herein). Specimens examined. – CANADA. ALBERTA: Edmonton, Emily Murphy Park, near LRT
bridge and Kinsmen Sports Centre, 53.52742, -113.514617, 2020, on moist N-facing mineral soil slope toe,
along trail, D. Haughland 2020-35 & P. Williams (hb. Haughland), D. Haughland 2020-37B & P. Williams
(hb. Haughland); Edmonton, Wolf Willow Creek, 53.502638, -113.602538, 2020, terricolous, D.
Haughland 2020-8 & C. Shier (hb. Haughland); Edmonton, Wolf Willow Creek, 53.50273, -113.602058,
2020, on moss, D. Haughland 2020-1 & C. Shier (hb. Haughland); Edmonton, Patricia Ravine, 53.504196,
-113.594313, 2020, on downed wood, D. Haughland 2020-58 & A. Hood (hb. Haughland); Edmonton,
River Loop Trail S of Fort Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland
2021-16 & S. Toni (hb. Haughland).
126
Figure 46. Tomentose Peltigera of Edmonton, plate 2 of 2: Peltigera praetextata. A, Typical habit with
ruffled lobes and tomentum limited to tips, Haughland 2020-58. B, Broad-lobed form resembling P.
membranacea, Haughland 2020-1. C, Lobules along margins and cracks, Haughland 2021-16. D, Lower
surface with raised pinkish-beige veins and simple, discrete rhizines, Haughland 2020-58.
Peltigera rufescens (Weiss) Humb. FIGURE 42 E-F.
River valley xylicole/terricole. Characterized by a tomentose to scabrid upper surface, typically
with small, grey to brown concave lobes with upturned lobe tips. The lower surface is characterised by low,
distinct veins, typically darkening centrally, and abundant rhizines, becoming enmeshed and hedgerow-like
centrally. Some specimens are sparsely lobulate, particularly along cracks. Molecular support: while
relatively poor, the single ITS sequence (isolate DLH19 from Haughland 2020-38) corresponds to
Peltigera rufescens 1, including containing the hypervariable region from Magain et al. (2018; Fig. 7
herein). Specimen examined. – CANADA. ALBERTA: Edmonton, Emily Murphy Park, near LRT bridge
and Kinsmen Sports Centre, 53.52742, -113.514617, 2020, on moist N-facing mineral soil slope toe, along
trail, D. Haughland 2020-38 & P. Williams (hb. Haughland); Edmonton, River Loop Trail S of Fort
Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland 2021-17 & S. Toni (hb.
Haughland).
127
Figure 47. Photobionts from the primary thalli of Chaenotheca species found in Edmonton. B is from a
collection outside of Edmonton. A, Filamentous, relatively small-celled alga Stichococcus from C.
trichialis, Haughland 2020-11B. B, Thick-walled Trentepohlia from C. hispidula, ca. 64 km NW of
Notikewin, ABMI Site 429, 57.54, -117.80, 2012, on bark, A. Hillman s.n. [ABMI Lichen # 120959].
GROUP 13. CALICIOID LICHENS AND FUNGI
Nine species. Key literature: McMullin et al. 2018; Selva 2013, 2014; Stordeur et al. 2013; Tibell
1996, 1999. The key below focuses on macroscopic features and niche; definitive identification requires
microscopic examination including identification of the photobiont through examination of the primary
thallus for lichenized species, and consultation with the resources listed. The two photobionts present in
species documented to date are illustrated here (Fig. 47).
1a. Ascomata completely black or brown, lacking pruina; no primary thallus visible; epiphytic on various
deciduous trees and shrubs; non-lichenized .................................................................................................... 2
1b. Ascomata with pruina on stalk or capitulum; primary thallus often visible on substrate; often associated
with Picea glauca in moist habitats; lichenized .............................................................................................. 6
2a. Stalk swelling in the middle; growing on creviced Populus bark ............................ Caliciopsis calicioides
2b. Stalk cylindrical, capitulum wider than stalk; host plant various .............................................................. 3
3a. Growing on Populus; ascomata 0.5–1 mm tall .......................................... Phaeocalicium populneum s.l.
3b. Growing on Betula, Alnus, or Rosa; ascomata typically <0.6 mm tall ...................................................... 4
4a. Growing on Alnus; at least some ascomata branching................................................ Stenocybe pullatula
4b. Growing on Betula or Rosa; no ascomata branching ................................................................................ 5
5a. Growing on Betula; stalk paler than capitulum; capitulum:stalk ratio approximately 1:3 ..........................
........................................................................................................................ Phaeocalicium aff. tremulicola
5b. Growing on Rosa; stalk concolorous with capitulum; capitulum:stalk ratio approximately 1:1 .................
................................................................................................................................. Phaeocalicium sp. nov. 1
6a. Primary thallus pale green, lime-green to yellow, farinose to granular ..................................................... 7
6b. Primary thallus not apparent or squamulose .............................................................................................. 8
7a. Stalks often >1mm tall and with yellowish-green pruina; capitula globose; growing on sheltered soil,
roots and wood on tree bases in riparian and ravine habitats ................................... Chaenotheca furfuracea
7b. Stalks typically <1 mm tall with light brown pruina; capitula lenticular (shaped like a lens when viewed
from the side); growing on the bases of Picea glauca, sometimes intermingled with Chaenotheca
furfuracea .................................................................................................................... Chaenotheca stemonea
8a. Primary thallus composed of grey-green waxy squamules; ascomata typically with white pruina .............
......................................................................................................................................Chaenotheca trichialis
8b. Primary thallus immersed/invisible; capitula with yellow pruina ......................... Chaenotheca hispidula
128
Figure 48. Caliciopsis calicioides, on Populus balsamifera, UoA-CC-90.
†Caliciopsis calicioides (Fr.) Fitzp. FIGURE 48.
Parkland epiphyte. This distinctive black calicioid has a central swelling that constitutes the
ascogenous region; distally a long narrow beak forms through which the spores are released. It can be
found in mature forests in the bark crevices of large-diameter Populus balsamifera. It is not clear why it is
often excluded from treatments of calicioid lichens and fungi. It is undoubtedly more common than existing
reports suggest. We are including it in part due to research suggesting it may be an indicator of healthy,
mature forests (Jordal et al. 2014). See Jordal et al. (2014) for additional morphological and habitat images.
Chemistry: not investigated. Molecular support: none found, one sequence in GenBank, no new sequences
generated.
Specimen examined & observations. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site
148E, 53.587522, -113.640605, 2019, on trunk of Populus balsamifera, D. Haughland & L. Hjartarson s.n.
[UoA-CC-90] (hb. Haughland); Edmonton, Urban Monitoring Site 163E, 53.6080134, -113.590864, 2019,
on trunk of Populus balsamifera, D. Haughland & L. Hjartarson (unvouchered observation).
Chaenotheca furfuracea (L.) Tibell FIGURE 49 A.
River valley terricole/basal epiphyte. This is a distinctive species that can be sighted from a
distance due to its almost glowing, yellow-green thallus. The dust-like primary thallus produces relatively
tall (0.5–1[–2] mm) yellow-pruinose stalks and globose capitula. It is by far the most common lichenized
calicioid in Edmonton, and can be found reliably at the bases of Picea in river valley parks, where the bole
and roots meet moist soil, in shaded nooks. The photobiont is Stichococcus, a small, filamentous,
129
Figure 49. Chaenotheca of Edmonton. A, C. furfuracea, Haughland 2020-11B. B, C. hispidula,
Haughland 2020-102. C, C. stemonea, Haughland 2020-11A. D, C. trichialis, Haughland 2019-117.
bacilliform green alga. Spores are spherical, yellow, to 3 µm in diameter. Care should be taken to check
colonies for co-occurring C. stemonea, which has brown-grey pruina, lenticular capitula and a PD+ orange
primary thallus. Chemistry: all spot tests negative, vulpinic acid, pulvinic acid, pulvinic dilactone (Tibell
1999). Molecular support: high species-level support albeit with limited sampling, reciprocally
monophyletic and sister to C. brachypoda (Ach.) Tibell (Tibell et al. 2019). No new sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf
Willow Creek, 53.502511, -113.601928, 2020, on bark on sheltered large roots of Picea glauca snag, D.
Haughland 2020-11B & C. Shier (hb. Haughland); Edmonton, Patricia Ravine, 53.503283, -113.5932,
2020, on base of mostly decorticate stump, on roots and bark and soil, D. Haughland 2020-40 & A. Hood
(hb. Haughland); Edmonton, Kinnaird Ravine, 53.558778, -113.462953, 2020, on base of Picea glauca, on
roots and bark and soil, D. Haughland 2020-26 & P. Williams (hb. Haughland); Edmonton, Hawrelak Park,
53.52, -113.54, 2012, on sheltered soil at base of tree, D. Haughland 2012-393 (hb. Haughland);
Edmonton, Larch Sanctuary, 53.449813, -113.551919, 2020, on base of mature Betula tree, D. Haughland
(unvouchered observation: NatureLynx record); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315,
2021, on decayed stump, D. Haughland (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/17450/details).
Chaenotheca hispidula (Ach.) Zahlbr. FIGURE 49 B.
River valley epiphyte. Apparently rare in Edmonton, this diminutive epiphyte is diagnosed by
short ascomata (~0.5 mm), yellow pruina on the upper stalk and capitulum, an association with
Trentepohlia, and globose spores 6–7 µm in diameter. Spore size should be checked to definitively exclude
130
C. olivaceorufa Vain., a similar species not yet detected in Alberta, with spores 3.0–4.5 µm in diameter
(McMullin et al. 2018). Chemistry: all spot tests negative, vulpinic acid in pruina (Tibell 1999). Molecular
support: not yet assessed at species-level, a single sequence basal to the Chaenotheca clade (Tibell 2001),
no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Kinnaird Ravine, 53.558953, -
113.459253, 2020, on bark of bole of Picea glauca snag, D. Haughland 2020-102 (hb. Haughland).
Chaenotheca stemonea (Ach.) Müll. Arg. FIGURE 49 C.
River valley epiphyte. This species may be uncommon or under-detected. It resembles
Chaenotheca furfuracea but has shorter stalks that are brown-grey-pruinose (vs. yellow-green in C.
furfuracea) and lenticular capita (vs. globose in C. furfuracea). As with C. furfuracea, the photobiont is
Stichococcus, a filamentous, bacilliform green algae (Fig. 47A). Spores are spherical, yellow, to 3–4 µm in
diameter. Chemistry: PD+ yellow to reddish, K-, KC-, C-, barbatic and obtusatic acid (Tibell 1999).
Molecular support: two sequences in GenBank, not assessed at species-level, no new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.502511, -
113.601928, 2020, on bark on sheltered large roots of Picea glauca snag, D. Haughland 2020-11A & C.
Shier (hb. Haughland).
Chaenotheca trichialis (Ach.) Th. Fr. FIGURE 49 D.
River valley epiphyte. Rare in Edmonton, this species typically has a unique waxy, blue-green
verrucose to squamulose thallus, producing shiny, black-stalked ascomata. The lower excipulum and
sometimes the upper stalk may be white-pruinose. The photobiont is Stichococcus (Fig. 47A). Spores
yellow, globose, up to 5.5 µm in diameter, slightly larger than indicated in the literature (Tibell 1999).
Chemistry: all spot tests negative, two unidentified substances (Middelborg & Mattsson 1987). Molecular
support: weak at species-level, paraphyletic with Chaenotheca xyloxena Nádv. (Tibell et al. 2019). No new
sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.50315, -
113.602186, 2019, on bark on sheltered large roots of Picea glauca snag, D. Haughland et al. 2019-117
(hb. Haughland); Edmonton, Kinnaird Ravine, 53.558953, -113.459253, 2020, on trunk of Picea glauca
snag, D. Haughland 2020-101 (hb. Haughland).
†Phaeocalicium populneum Brond. ex Duby s.l. FIGURE 50 A.
River valley and parkland epiphyte. Common on Populus across Alberta, from saplings to mature
trees. This species is characterized by brown to black stalks bearing cupulate to lenticular capitula,
periclinal hyphae forming the excipulum and uniseptate, brown spores with walls that range from smooth
to minutely areolate in ornamentation at 1000x. Across Alberta, this species is highly variable in spore size,
ascoma height and even capitulum shape, and may include multiple, undescribed species (Haughland et al,
in prep.). Edmonton material: ascomata to 0.5–1.0 mm tall, black globose to lenticular capitulum, rarely
branching. The spores are brown, uniseptate, septum pigmented and slightly constricted or wavy, spore
wall smooth to faintly areolate ornamented, occasionally becoming 3-septate, 13–16 × 4.5–5.5 µm (only
typical uniseptate spores measured). Chemistry: all Edmonton collections had purple or blackish pigments
in the lower stalk that turned K+ aeruginose green, either in small patches or the entire lower stalk.
Otherwise, the stalk and capitulum are red-brown in squash, swelling but K-. The collection from the rural
Ardrossan Air Quality Monitoring Station was the exception, and it was both more robust (similar to much
of Alberta material from non-urban settings) and K-. Substances detected by TLC: not investigated. The
literature is mixed on chemistry; Tibell (1996, 1999) and Selva (2014) indicate K- while Aguirre-Hudson &
Spooner (2019) and Nimis (2016) report K+green reaction in the stalk. Molecular support: additional
molecular work is needed to determine if the urban phenotype is in response to growth in an urban
environment or indicative of a different genotype. Species-level support for Phaeocalicium populneum s.s.
has not been assessed phylogenetically, two sequences in GenBank, sequencing in progress with S. Selva
and T. McMulllin.
131
Figure 50. Non-lichenized calicioid fungi of Edmonton. A, Phaeocalicium populneum s.l., Haughland
2013-247. B, Phaeocalicium aff. tremulicola, Haughland 2013-192. C, Phaeocalicium sp. nov. 1,
Haughland 2017-376. D, Stenocybe pullatula, Haughland 2020-15B.
Representative specimens examined. – CANADA. ALBERTA: Ardrossan, Air Quality
Monitoring Station, 53.55468, -113.143238, 2019, on trunk of Populus balsamifera, D. Thauvette & J.
Birch s.n. [UoA-CC-89] (hb. Haughland); Edmonton, Whitemud Creek, 53.50, -113.56, 2013, on Populus
branches, D. Haughland et al. 2013-247 (hb. Haughland); Edmonton, Whitemud Creek, 53.501458, -
113.560761, 2020, on Populus balsamifera dead branch, D. Haughland 2020-62 & P. Williams (hb.
Haughland); Edmonton, Buena Vista Meadow off-leash park, 53.517138, -113.549195, 2020, on Populus
tremuloides dead vertical branch, D. Haughland 2020-92 & P. Williams (hb. Haughland); Edmonton,
Buena Vista Meadow off-leash park, 53.519725, -113.545738, 2020, on Populus tremuloides live
horizontal branch, D. Haughland 2020-93 & P. Williams (hb. Haughland); Edmonton, Sir Wilfrid Laurier
Park, 53.50834, -113.560926, 2019, on bark of large downed Populus, D. Haughland 2019-115A & P.
Williams (hb. Haughland); Edmonton, MacKenzie Ravine, 1976, on tree bark, D.C. Lindsay s.n. (PMAE-
B77.24.170).
***†Phaeocalicium aff. tremulicola (Norrlin ex Nyl.) Tibell FIGURE 50 B.
River valley epiphyte. A single collection was made of a diminutive Phaeocalicium on live Betula
twigs. It is closest to Phaeocalicium tremulicola (Norrlin ex Nyl.) Tibell anatomically, however, P.
tremulicola has only been documented on Populus tremula (Tibell 1999) and Hamamelis virginiana (Selva
2014). Edmonton material: ascomata 0.3–0.4 mm tall, pale brown, with globose to obovate capitula. The
132
stalks are pale pink-brown in squash to almost hyaline at the base with an outer hyaline layer, and the
excipulum has an outer layer of thick-walled isodiametric cells. The spores are brown, smooth, broadly
fusiform, with tips pointed to mucronate; most commonly they are uniseptate, occasionally becoming 3-
septate, slightly constricted at the well-pigmented septa, 12–15 × 5–6 µm. Chemistry: not investigated.
Molecular support: in progress, S. Selva has examined the material and noted “not P. tremulicola, possibly
a new species” (S. Selva, pers. comm. 2020). No sequences of P. tremulicola s.s. in GenBank, sequencing
with S. Selva and T. McMullin is in progress.
Specimen examined. – CANADA. ALBERTA: Edmonton, Whitemud Ravine, 53.50, -113.56,
2013, on live Betula papyrifera twig, D. Haughland et al. 2013-192 (hb. Haughland).
***†Phaeocalicium sp. nov. 1, ined. Haughland FIGURE 50 C.
River valley and parkland epiphyte. This undescribed species was first found on dead Rosa
branches in the Parkland Natural Region at an ABMI site. Searches by the senior author have found it to be
common on relatively tall, dead (rarely live) Rosa branches in at least partially protected habitats in the
river valley parks of Edmonton. Host Rosa that could be identified to species were R. woodsii. To date we
do not know of any populations outside of Alberta, despite searches by the senior author in Saskatchewan,
Yukon Territory, and the Northwest Territories in similar habitat. To our knowledge this is the first
calicioid found on Rosa. We provide a basic description to alert others to this unique calicioid so that
additional populations may be sought. Edmonton material: dark brown ascomata often growing near branch
nodes, to 0.4 mm tall with the cupulate capitulum typically comprising ≥50% of the height. Development
of the capitulum starts before stalk elongation. The stalk has an outer hyaline coat over isodiametric cells
that continue upwards to form the outer excipulum, becoming stretched horizontally around upper edge of
the capitulum. Asci are cylindrical, 75–88 µm long, spores brown, 10–15 × 4.5–6 µm, mostly uniseptate,
some spores forming two additional septa even within the ascus, spore wall with faint, minute areolate
ornamentation visible at 1000x. While most spores remain within the asci, loose spores are visible and may
form a tall mazaedial mass in some stalks. Chemistry: not investigated. Molecular support: sequencing is in
progress to confirm the phylogenetic placement of this species with T. McMullin and S. Selva.
Representative specimens examined. – CANADA. ALBERTA: Edmonton, Terwillegar Park,
53.4788, -113.6218, 2017, on dead standing stem of Rosa cf. woodsii, D. Haughland 2017-375 & P.
Williams (hb. Haughland); Edmonton, Buena Vista Meadow, 53.521469, -113.548092, 2017, on dead
standing stem of Rosa woodsii, D. Haughland 2017-376 (hb. Haughland); Edmonton, Buena Vista
Meadow, 53.521127, -113.548707, 2017, on live stem of Rosa woodsii, D. Haughland 2017-377 & D.
Thauvette (hb. Haughland).
†Stenocybe pullatula (Ach.) Stein FIGURE 50 D.
River valley epiphyte. Common and variable calicioid fungus on Alnus, characterized by short
ascomata (typically <0.6 mm tall but in some material in Alberta exceeding 1 mm), branching, with
narrow, vertically striate capitula. Spores average 15–20 × 4.0–5.5 µm, brown, smooth to minutely
ornamented, fusiform (with pointed tips), 1–3-septate, septa typically poorly pigmented but occasionally
dark and constricted. Edmonton material fits within the range of variation observed across Alberta.
Chemistry: all spot tests negative, no secondary metabolites detected (Tibell 1999). Molecular support:
genus and species-level not assessed phylogenetically, three sequences in GenBank, a single sequence
forms a highly supported branch sister to single sequence of Phaeocalicium populneum (Tibell & Vinuesa
2005). Specimens examined. – CANADA. ALBERTA: Edmonton, Whitemud Ravine, 53.491661, -
113.55914, 2020, on bark of dead Alnus snag, D. Haughland 2020-15B & P. Williams (hb. Haughland);
Edmonton, Whitemud Park, 53.50, -113.56, 2013, on Alnus branches, D. Haughland 2013-248 & S. Selva
(hb. Haughland); Edmonton, Whitemud Park, 53.501717, -113.560812, 2020, on Alnus incana subsp.
tenuifolia branches, D. Haughland 2020-63 & P. Williams (hb. Haughland).
133
GROUP 14. CLADONIA
Fifteen species. Key literature: Ahti & Stenroos 2013; Brodo et al. 2001; Goward 1999. A diverse
genus of over 90 species in Alberta, Cladonia is underrepresented in Edmonton in both diversity and
biomass. Given these are fairly detectable species, we suspect the dearth of Cladonia in our urban
environment is real; while additional species are likely to be found (including perhaps C. rei Schaerer,
Haughland et al. 2018), the depauperate flora documented to date is reminiscent of poor boreal sites. We
employ a slight simplification of the morphogroup system of Goward (1999), while maintaining Goward’s
original groups to aid the user in working between the more inclusive keys to species of British Columbia
(Goward 1999) and Alberta (Haughland, unpublished), and these species-limited keys. Spot tests are
helpful for identification, particularly for novices learning to differentiate the pale yellow of usnic acid-
producing species. This is a difficult group both morphologically and phylogenetically, with many species
as currently circumscribed polyphyletic with similar species (e.g., Pino-Bodas et al. 2011, 2015; Stenroos et
al. 2002, 2018). Expect future taxonomic revisions.
Key to the Cladonia Groups
1a. Podetia absent; thallus consisting entirely of primary squamules. In Edmonton, this group is composed
of immature thalli that cannot be identified to species using morphology or chemistry ...................................
................................................................................................... Group A - Cladonia spp., not treated further
1b. Podetia present; primary thallus either crustose or squamulose or missing .............................................. 2
2a. Podetia richly branched (more than two times) ............................................................................ Group B
2b. Podetia unbranched to slightly branched (once or twice) ......................................................................... 3
3a. Some podetia tips opening by a gaping hole, look for in-rolled margins around the hole as confirmation
that the hole developed as part of the podetium (vs. a broken podetium) .......................................... Group C
3b. Podetia tips closed ..................................................................................................................................... 4
4a. Podetia solid/compact throughout, longitudinally ribbed or fibrous AND esorediate AND cupless ..........
............................................................................................................................................................ Group D
4b. Podetia not longitudinally ribbed or fibrous, stalk hollow and tubular, may be sorediate or cupped ....... 5
5a. Podetia distinctly cupped, cups typically symmetrical and wider than the stalk ....................................... 6
5b. Podetia wand-like to antler-like, typically either lacking cups, cups shallow and no wider than the stalk,
or with sparse, irregular, oblique cups ............................................................................................................. 8
6a. Cortex with a distinct yellowish cast (usnic acid present); apothecia red or less commonly beige to
brown .................................................. Group E – None known from Edmonton at present, not treated further
6b. Cortex not distinctly yellowish (usnic acid absent or in low concentration); apothecia never red ............ 7
7a. Podetia bearing at least some soredia (check upper portions) ...................................................... Group F
7b. Podetia lacking soredia ................................................................................................................ Group H
8a. Podetia bearing soredia or corticate granules, or both (check upper portions) .............................. Group I
8b. Podetia lacking soredia and corticate granules (note: dorsiventral squamules and/or microsquamules
may be present) .................................................................................................................................. Group J
Cladonia Group B: branched > 2 times
1a. Podetia lacking a cortex: podetial surface appressed-fibrous under the microscope (the “reindeer”
lichens) ......................................................................................................... Cladonia arbuscula subsp. mitis
1b. Podetia corticate at least in part – matte or shiny, but fibrous hyphae not visible under the microscope . 2
134
2a. Podetia tips and/or branch axils bearing flaring cups, these closed but sieve-like, doily-like, or lacerate
OR basal portions of main podetia more or less distinctly flattened in cross-section; terminal portions of the
supporting branches somewhat flattened or longitudinally lacerate, bearing abundant apothecia at the tips ....
....................................................................................................................................... Cladonia multiformis
2b. Podetia uncupped, terminal portions of branches with narrow openings or lacerations into the interior,
these restricted to branch axils, not at all sieve-like; terminal portions of podetia in part lacking a cortex,
often with soredia, corticate granules and/or detachable microsquamules, these sometimes extending
downward into the middle or basal portions of the podetia .......................................... Cladonia scabriuscula
Cladonia Group C: podetia tips with open axils or open cups
1a. Largest apical cups open and distinctly flaring; podetia in Edmonton stout, seldom branching; UV+
blue-white, PD- .............................................................................................. Cladonia crispata var. crispata
1b. Apices open but not flaring OR cups with seive-like perforations; podetia in Edmonton relatively tall,
often branching; UV-, PD+ red to orange ....................................................... Go to Group B Key couplet 2
Cladonia Group D: podetia longitudinally ribbed or fibrous
1a. Apothecia beige or red; common on lignum and downed logs; podetia with a distinctly yellowish cast
(usnic acid), K-, KC+ yellow .......................................................................................................................... 2
1b. Apothecia dark brown; on soil, mossy rock or anthropogenic substrates; podetia grey, not at all
yellowish, K+ yellow or red, KC- ......................................................................................... Cladonia cariosa
2a. Apothecia beige to pale brown ....................................................................................... Cladonia botrytes
2b. Apothecia red .............................................................................................................. Cladonia cristatella
Cladonia Group F: podetia cupped, sorediate, grey, green or brown
1a. Podetia more green than grey, not melanotic (blackening); soredia fine or coarse; PD+ orange to red,
UV- .................................................................................................................................................................. 2
1b. Podetia greyish-white to grey-green, often partially melanotic especially at the base; soredia granular;
PD- or PD+ orange to red, UV+ purple to white; not yet found within Edmonton, but confirmed locations
close to study boundary .....................................................................................................................................
.................................. Cladonia spp. including C. grayi G. Merr. ex Sandst. & C. merochlorophaea Asahina
see discussion under Cladonia chlorophaea
2a. Podetia with floury, fine soredia, >1 layer deep covering stalk; outer and inner cup, cups typically
abruptly flaring with clear delimitation between cup and stalk ......................................... Cladonia fimbriata
2b. Podetia with granular, coarse soredia, typically patchy or sparse; cup flaring gradually from the
podetium ....................................................................................................................... Cladonia chlorophaea
Cladonia Group H: podetia cupped, esorediate
1a. Podetia commonly >1.5 cm tall, variable but often with relatively large, bulbous, brown apothecia on
the margins of the cups, secondary squamules absent or limited to the stalk, inside of the cup smoothly
corticate ..................................................................................................... Cladonia gracilis subsp. turbinata
1b. Podetia typically <1.5 cm tall, squat, with peltate, pancake-like squamules inside the cup ........................
............................................................................................................................................. Cladonia pyxidata
Cladonia Group I: sorediate podetia lacking regular cups
1a. Podetial soredia continuous, not at all borne in discrete patches; color consistent throughout podetium ...
......................................................................................................................................................................... 2
1b. Podetial soredia discontinuous, at least in part borne in discrete patches; podetia often mottled green
and white ......................................................................................................................................................... 4
135
2a. Podetia green, wand-like (tapered from bottom to top); primary squamules present and conspicuous;
cortex of the podetium often intact and esorediate basally; PD+ orange to red .............. Cladonia coniocraea
2b. Podetia pale mint-green to grey-green, often with swollen, club-shaped or antler-shaped tips; primary
squamules various, but often small to absent; PD- or PD+ orange to red ....................................................... 3
3a. Podetia pale minty-green, often slightly swollen or club-shaped at the very tips, occasionally branched
near the terminus, often tipped with small red apothecia or pycnidia; PD-, UV- or UV+ faint white ..............
................................................................................................................... Cladonia macilenta var. bacillaris
3a. Podetia grey-green, typically sorediate over entire podetium, forming irregular, oblique cups or antler-
like proliferations, apothecia brown and pycnidia black but both very rare; PD+ orange to red, UV- ..............
............................................................................................................................................. Cladonia subulata
4a. Podetia <2 cm tall, variable from wand-like to terminating in small cups with narrow, pointed
proliferations like a jester-cap; most common chemotype is PD-, UV+ faint white or UV- ........ Cladonia rei
4b. Podetia commonly >2.5 cm tall, most of the cortex intact with clearly delimited patchy soralia towards
the tip; PD+ red to orange, UV- ................................................................. Cladonia cornuta subsp. cornuta
Cladonia Group J: podetia uncupped, esorediate
1a. Apothecia beige to pale brown ....................................................................................... Cladonia botrytes
1b. Apothecia red .............................................................................................................. Cladonia cristatella
Cladonia arbuscula subsp. mitis (Sandst.) Ruoss
(≡ Cladonia mitis Sandst., ≡ Cladina mitis (Sandst.) Mong.) FIGURE 51 D.
River valley and ravine system xylicole. This is the most common reindeer lichen in Alberta,
occurring on lignum, soil, and moss. In Edmonton it is rare and poorly developed, and has been found only
on lignum. Primary thallus crustose, evanescent, forming ecorticate podetia to 12 cm tall that are densely
branched. The podetia typically are yellowish-grey, with an almost transparent stereome at the base. The
tips branch 3–4 times, and often form either brown apothecia or pycnidia. Chemistry: K-, KC+ yellow, C-,
UV254-, PD-. Substances detected by TLC: usnic acid, isousnic acid, ±rangiformic acid (trace). Molecular
support: still incompletely understood at species-level (Ahti & Stenroos 2013), forming a monophyletic
clade nested within Cladonia arbuscula (Wallr.) Flot. (Piercey-Normore et al. 2010).
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
53.502711, -113.60241, 2020, lignicolous on downed log, D. Haughland 2020-10B & C. Shier (hb.
Haughland); Edmonton, W of Whitemud Park, along Grandview Stairs, 53.502357, -113.552249, 2020, on
lignum, L. Hjartarson (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/12334/details).
Cladonia botrytes (K. G. Hagen) Willd. FIGURE 51 A.
River valley and ravine system xylicole. Primary thallus squamulose, developing pale yellow,
esorediate podetia to 2(–3) cm tall that can be moderately branched in upper parts. The podetia always
terminate in ochraceous to pale-brown apothecia. This is a common boreal species that may be tolerant of
moderately polluted environments (Ahti & Stenroos 2013). Chemistry: K-, KC+ yellow, PD-, UV254/365-.
Secondary metabolites detected by TLC: usnic acid, barbatic acid. Molecular support: strong, multiple
sequences cluster in a well-supported branch in the Clade “Ochroleucae” (Stenroos et al. 2018). No new
sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton,
Patricia Ravine, 53.502112, -113.592209, 2020, on downed wood, S. Toni (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/12143/details); Edmonton, MacKenzie Ravine,
53.529146, -113.558821, 2020, on downed log, D. Haughland (unvouchered observation: NatureLynx
record https://naturelynx.ca/sightings/14205/details); Edmonton, near Northland sandpit, 2 mi W and 1 mi
S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.101); Edmonton, MacKenzie
Ravine, 1976, on rotten log, D.C. Lindsay s.n. (PMAE-B77.24.163); Edmonton, MacKenzie Ravine
boardwalk, 53.528875, -113.558827, 2021, on decayed Picea glauca log, D. Haughland 2021-8 (hb.
Haughland).
136
Figure 51. Fibrous and shrubby Cladonia of Edmonton. A, C. botrytes, growing on lignin of large log in
river valley, Haughland 2021-8. B, C. cariosa, Dueck https://naturelynx.ca/sightings/9803/details. C, C.
cristatella, Morinville region north of Edmonton, Fielder https://naturelynx.ca/sightings/15528/details. D,
C. arbuscula subsp. mitis, Hjartarson https://naturelynx.ca/sightings/12334/details.
Cladonia cariosa (Ach.) Sprengel FIGURE 51 B.
Apparently rare ravine terricole. Primary squamules tend to be small, to 2–3 mm across, convex
(like clamshells), and persistent. When podetia are absent, confident identification without genetics is not
possible because of overlapping chemistry with immature squamules of other Cladonia. Podetia to 3 cm
tall, slender, made of solid, grey to grey-green, fissured cartilaginous strands, branching distally and always
bearing brown apothecia. No vegetative propagules present. This species apparently can thrive in
anthropogenically altered habitats, commonly growing on disturbed soil. Similar species include Cladonia
symphycarpa (Flörke) Fr. and related, currently undescribed lineages (Lewis 2022: podetia may be similar,
but primary squamules are larger and thicker, growing to 1 cm long, often reflexed when dry; chemistry
may include psoromic or norstictic acids) and C. acuminata (Ach.) Norrlin (primary squamules similar but
podetia wand-like, not fibrous, typically unbranched, lacking apothecia, chemistry atranorin ± norstictic,
connorstictic acids). Chemistry: PD+ yellow, orange or red, K- or K+ dingy brown or yellow, KC-, UV-.
Secondary metabolites detected by TLC: atranorin, ±fumarprotocetraric acid, protocetraric acid,
rangiformic acid, and norrangiformic acid. Molecular support: a genotyping-by-sequencing study provides
strong support for this species (Lewis 2022). Previous phylogenetic work using multiple loci left some
phenotypically-similar lineages unresolved (Pino-Bodas et al. 2012).
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Mill Creek Ravine
South, 53.509826, -113.463148, 2019, on landscaping fabric over rocks, T.L. Dueck (unvouchered
137
observation: NatureLynx record https://naturelynx.ca/sightings/9803/details); Edmonton, Mill Creek
Ravine South, 53.511504, -113.465939, 2020, terricolous, T.L. Dueck (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/13503/details); Edmonton, River Loop Trail S of Fort
Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland 2021-19 & S. Toni (hb.
Haughland, NatureLynx record https://naturelynx.ca/sightings/17285/details).
Cladonia chlorophaea (Flörke ex Sommerf.) Sprengel FIGURE 52 A-B.
River valley and ravine system xylicole and epiphyte. Primary thallus squamulose, podetia to 4 cm
tall, characterized by broad green to blue-green cups with granular soredia in the upper portion,
subcontinuously corticate towards the base. Apothecia are fairly common, on short marginal proliferations.
The Cladonia chlorophaea group generally requires chemistry to identify with certainty, although with
experience color can discriminate this species from the more acid-tolerant and typically paler C. grayi.
(with grayanic acid and ±fumarprotocetraric acid, PD+ orange or PD-, UV254+ violet). and C.
merochlorophaea (with merochlorophaeic acid, 4-O-methylcryptochlorophaeic acid, cryptochlorophaeic
acid and ±fumarprotocetraric acid, PD+ orange or PD-, UV254+ blue-white). Even with chemistry,
identification remains challenging. Morphology intergrades with the trumpet-shaped, farinose-sorediate C.
fimbriata and the esorediate C. pyxidata, and the chemistry of these three is identical. Chemistry: K- or K+
dingy brown, KC-, C-, PD+ orange, UV254-. Secondary metabolites detected by TLC: fumarprotocetraric
acid. Molecular support: unresolved within the Clade “Cladonia”, subclade “Graciles” (Stenroos et al.
2018), under further investigation (Ahti & Stenroos 2013). No new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Hawrelak Trail off-leash park,
53.520733, -113.54158, 2020, on Betula base, D. Haughland 2020-19A (hb. Haughland); Edmonton, near
Northland sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-
B77.24.105); Edmonton, Whitemud Park, 1976, on wood, D.C. Lindsay s.n. (PMAE-B77.24.121);
Edmonton, MacKenzie Ravine, 1976, on rotting log, D.C. Lindsay s.n. (PMAE-B77.24.186); Edmonton,
MacKenzie Ravine, 53.52904, -113.5605, 2020, on stump on edge of mineral seep, D. Haughland 2020-
107 (hb. Haughland, NatureLynx record https://naturelynx.ca/sightings/14204/details).
Cladonia coniocraea (Flörke) Sprengel FIGURE 53 A.
River valley and ravine system xylicole and epiphyte. A variable, wand-like Cladonia with a
persistent primary thallus of green squamules. The podetia are 1–3 cm tall, mint-green to grey-green,
unbranched and slender, often tapering to the tip, typically continuously sorediate except near the podetial
base where the cortex remains intact. Apothecia are infrequent, brown, but may be found in very mature
specimens that develop narrow cups at the podetial tips. This species can be separated from C. subulata by
the latter’s more continuous, often grey-brown soredia, typically evanescent primary squamules, and a
proclivity to form antler-like branches or proliferations from cups. Unfortunately, intermediates exist, and
chemistry does not distinguish them. Another species that may co-occur is C. rei, but in Alberta, C. rei and
is a grassy-green color (vs. mint-green), and almost always lacks furmarprotocetraric acid, instead
producing only homosekikaic acid ± sekikaic acid, (PD-, UV254- or +dull white; Haughland et al. 2018).
Chemistry: PD+ orange, UV254-, K- or K+ dingy brown, C-, KC-. Substances detected by TLC:
fumarprotocetraric acid, sometimes with traces of unknown compounds. Molecular support: strong species-
level support within the C. gracilis group if synonymy with C. ochrochlora Flörke and C. cornuta subsp.
groenlandica (E. Dahl) Ahti is accepted (Pino-Bodas et al. 2011, Stenroos et al. 2018). No new sequences
generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton
Hawrelak Park, 53.52, -113.54, 2017, on decaying wood under Betula papyrifera, D. Haughland et al.
(unvouchered observation); Edmonton, Whitemud Ravine, 53.499008, -113.560730, 2020, on tree base, D.
Haughland & P. Williams s.n. (unvouchered observation; NatureLynx record https://naturelynx.
ca/sightings/12608/details); Edmonton, Whitemud Park, 53.503845, -113.554541, 2020, on moss over
wood, D. Haughland s.n. (unvouchered observation; https://naturelynx.ca/sightings/13871/details);
Edmonton, 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.104).
138
Figure 52. Podetia and colonies of short sorediate or squamulose cupped Cladonia of Edmonton. A-B, C.
chlorophaea, growing on stump, with sparse, granular soredia, Haughland 2020-107. C-D, C. fimbriata. C,
Cups with multiple layers of finer-grained soredia, Haughland 2021-24. D, Colony on base of Betula,
Haughland https://naturelynx.ca/sightings/12605/details. E-F, C. pyxidata, growing on base of Betula, with
peltate squamules in the cup, Haughland 2020-19B.
139
Figure 53. Wand-like Cladonia of Edmonton. A, C. coniocraea, Haughland https://naturelynx.ca/
sightings/13871/details. B, C. cornuta subsp. cornuta, Fielder https://naturelynx.ca/sightings/16159/details.
C, C. macilenta var. bacillaris, Toni https://naturelynx.ca/sightings/12140/details. D, C. scabriuscula,
Haughland https://naturelynx.ca/sightings/10685/details. E, C. subulata, Haughland 2021-15.
140
Cladonia cornuta (L.) Hoffm. subsp. cornuta FIGURE 53 B.
River valley and ravine system terricole and xylicole. A common and often prolific wand-like
Cladonia with a persistent to more often evanescent primary thallus. This species forms tall (to 8 cm)
unbranched podetia with round patchy soralia in the upper third. Podetia vary from green to brown to
spotted (shade-form?) due to a patchy algal layer. Chemistry: PD+ orange, UV254-, K- or K+ dingy brown,
C-, KC-. Substances detected by TLC: fumarprotocetraric acid, sometimes with traces of unknown
compounds. Molecular support: good species-level support based on limited sequences within the Clade
“Cladonia”, subclade “Graciles” (Pino-Bodas et al. 2011, Stenroos et al. 2018). No new sequences
generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
2020, lignicolous on downed log, 53.502711, -113.60241, D. Haughland & C. Shier s.n. (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/14347/details); Edmonton, near Northland
sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.97);
Edmonton, MacKenzie Ravine, 1976, on rotting log, D.C. Lindsay s.n. (PMAE-B77.24.184).
Cladonia crispata (Ach.) Flot. var. crispata FIGURE 54 A-B.
River valley and ravine system xylicole. Primary thallus squamulose, often with barrel-shaped
pycnidia forming laminally on the squamules. The open-cupped podetia are 2–6 cm tall, esorediate, and
often proliferate from the edges. Most diagnostic however are the UV+ open gaping funnels and cups, often
with dentate, pycnidiate margins. Chemistry: K-, KC-, C-, UV254+ white, PD-. Secondary metabolites
detected by TLC: squamatic acid. Molecular support: species-level support low, polyphyletic as currently
phenotypically delimited (Stenroos et al. 2018). No new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.502711, -
113.60241, 2020, lignicolous on downed log, D. Haughland 2020-10C & C. Shier (hb. Haughland);
Edmonton, MacKenzie Ravine boardwalk, 53.528875, -113.558827, 2021, on decayed Picea glauca log, D.
Haughland 2021-4 (hb. Haughland).
Cladonia cristatella Tuck. FIGURE 51 C.
Occasional river valley and ravine system xylicole. Common in the boreal on lignum or soil,
forming predictable communities with Cladonia arbuscula subsp. mitis, C. botrytes, and C. crispata var.
crispata. Primary thallus squamulose, often bearing barrel-shaped pycnidia with red pycnidial jelly visible
at the ostiole. The pale-yellow podetia are uncupped and often sparsely branched, up to 2.5 cm tall,
esorediate, with a continuous cortex. The bright red apothecia (or arrangement of apothecia sometimes
giving the illusion of cups) combined with the lack of soredia or true cups are diagnostic. In other regions
of Alberta, a rare species that may overlap morphologically is C. bellidiflora (Ach.) Schaerer, but that
species is UV+ white due to the presence of squamatic acid. Chemistry: K-, KC+ yellow, PD-, UV254/365- or
UV254/365 + pale yellow-white. Secondary metabolites detected by TLC: usnic acid, barbatic acid, ±4-O-
demethylbarbatic acid, ±didymic acid, rhodocladonic acid in apothecia. Molecular support: not assessed at
species-level, a single sequence forming a clade with either C. metacorallifera Asahina (Stenroos et al.
2002) or C. camerunensis Ahti & Flakus (Stenroos et al. 2018). No new sequences generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
53.502711, -113.60241, 2020, lignicolous on downed log, D. Haughland & C. Shier (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/14346/details); Edmonton, near Northland
sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-as minor
component in B77.24.101), D.C. Lindsay s.n. (PMAE-as minor component in B77.24.100).
Cladonia fimbriata (L.) Fr. FIGURE 52 C-D.
Apparently rare river valley and ravine system xylicole. Primary thallus squamulose, persistent,
producing podetia 0.5–3.0 cm tall. The bright-green to almost usnic-green podetia form abruptly flaring
trumpet-shaped cups with margins that are entire to slightly dentate. The podetia are coated with farinose
(floury), fine-grained soredia (see Fig. 43C). See Cladonia chlorophaea entry for similar species and how
to discriminate them. An additional similar species recently detected in Alberta is C. conista (Nyl.) Robbins
141
(ABMI 2020). Cladonia conista is similar in shape, but it tends to be largely corticate with soredia only on
the upper edge and inside of the cups; it also can be discriminated with TLC (but not with spot tests) as it
contains the fatty acid bourgeanic acid in addition to fumarprotocetraric acid. Chemistry: K- or K+ dingy
brown, KC-, C-, UV254-, PD+ orange. Secondary metabolites detected by TLC: fumarprotocetraric acid.
Molecular support: not assessed at species-level, a single sequence formed a clade with C. subsquamosa
Kremp. in early analyses (Stenroos et al. 2002), while two sequences cluster with low support with C.
chlorophaea (Stenroos et al. 2018). No new sequences generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Hawrelak Park,
Alberta, 53.52, -113.54, 2017, on decaying wood under Betula papyrifera, D. Haughland et al.
s.n.(unvouchered observation); Edmonton, Emily Murphy Park, near LRT bridge and Kinsmen Sports
Centre, 53.527, -113.51442, 2020, epixylic on decayed wooden bridge edge, D. Haughland & P. Williams
(unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/13877/details); Edmonton,
Whitemud Park, 53.49903, -113.560691, 2020, on tree base, hanna1025 (unvouchered observation:
iNaturalist record https://www.inaturalist.org/observations/59075224); Edmonton, Terwillegar Footbridge,
53.4797, -113.594315, 2021, on decayed stump, D. Haughland 2021-24 (hb. Haughland).
Cladonia gracilis subsp. turbinata (Ach.) Ahti FIGURE 54 C-D.
River valley and ravine system terricole and xylicole. Primary thallus squamulose, evanescent or
persisting, producing podetia 2–5 cm tall. The podetia are esorediate with a continuous, often hard, shiny
cortex and are often secondarily squamulose (shade form). The podetia form well-developed closed cups,
topped with pycnidia and/or large brown apothecia on marginal proliferations. This is one of the most
common Cladonia species in Alberta, occurring in every natural region, typically on soil, moss, debris, or
downed wood (ABMI 2020). The occasional perforate cup can be discriminated from C. multiformis by the
rarity of perforations and the larger, bulbous apothecia that are common in this species. Other similar, PD+
orange species not known from Edmonton include C. phyllophora Hoffm. (more northern in distribution,
outer cortex matte, dull and appearing fibrous), and C. ecmocyna (known only from the Rocky Mountains
and Foothills in Alberta, grey to grey-green, often with pruina on the distal half of the podetia, lacking large
apothecia, and K+ yellow due to atranorin). Chemistry: K- or K+ dingy brown, KC-, C-, UV254-, PD+
orange. Secondary metabolites detected by TLC: fumarprotocetraric acid. Molecular support: unresolved
within the Clade “Cladonia”, subclade “Graciles”, polyphyletic as currently morphologically
circumscribed but separate from C. ecmocyna (Fontaine et al. 2010, Pino-Bodas et al. 2011, Stenroos et al.
2018). No new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.502711, -
113.60241, 2020, lignicolous on downed log, D. Haughland 2020-10A & C. Shier (hb. Haughland);
Edmonton, near Northland sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on soil, D.C. Lindsay
s.n. (PMAE-B77.24.192), D.C. Lindsay s.n. (PMAE-B77.24.100).
Cladonia macilenta var. bacillaris (Ach.) Schaerer FIGURE 53 C.
Ravine system xylicole. Another common Cladonia of Alberta forests, it is apparently rare within
Edmonton. The primary squamules typically are persistent and variable in size and attachment to the
substrate (appressed to ascending). Podetia 0.5–1.5 cm tall, pale greenish-grey to pale mint-green,
unbranched to slightly-branched close to the blunt tips, covered in abundantly farinose soredia and
commonly tipped with small, red apothecia. This species can be discriminated by chemistry if the red
apothecia are lacking: C. coniocraea is PD+ orange, and C. bacilliformis (Nyl.) Sarnth. is KC+ yellow
(usnic acid), both species have brown apothecia if fertile. Chemistry: there are two chemical variants of this
species and the only one found in Alberta to date is C. macilenta var. bacillaris, K-, KC+ pinkish-gold, C-,
UV254- or dull/faint white, PD-. Secondary metabolites detected by TLC: barbatic acid, ±4-O-
demethylbarbatic acid, ±didymic acid. Molecular support: unresolved as currently morphologically
circumscribed, forming a highly supported clade with C. floerkeana (Fr.) Flörke (Stenroos et al. 2018). No
new sequences generated.
Observation. – CANADA. ALBERTA: Edmonton, Patricia Ravine, 53.502112, -113.592209,
2020, on downed wood, S. Toni (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/12140/details); Edmonton, Rainbow Valley, 1961, G.W. Scotter s.n.
(ALTA); Edmonton, MacKenzie Ravine boardwalk, 53.528875, -113.558827, 2021, on decayed Picea
glauca log, D. Haughland 2021-9 (hb. Haughland).
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Figure 54. Podetia and colonies of cupped, esorediate Cladonia of Edmonton. A-B, C. crispata var.
crispata, Haughland 2021-4. C-D, C. gracilis subsp. turbinata, with a few podetia of C. crispata var.
crispata, Haughland 2020-10A. E, C. multiformis cups, Fielder https://naturelynx.ca/sightings/5231/details.
F, C. multiformis colony, Dueck https://naturelynx.ca/sightings/14146/details.
143
Cladonia multiformis G. Merr. FIGURE 54 E-F.
River valley and parkland terricole. A variable species that can have podetia that are either
branched and fertile with flattened branches and longitudinal perforations, or cupped with sieve-like or
doily-like perforations in the top and bottom of the cup. Both morphs are present in Edmonton and in
Alberta. Similar in size and chemistry to C. gracilis subsp. turbinata, the regular perforations and small
apothecia of C. multiformis help to discriminate it. Cladonia crispata var. crispata can also resemble this
species, but crispata is PD- and UV+ white. Chemistry: K- or K+ dingy brown, KC-, C-, UV254-, PD+
orange. Substances detected by TLC: fumarprotocetraric acid. Molecular support: unresolved and
polyphyletic, in a complex with C. farinacea (Vain.) A. Evans, C. furcata (Hudson) Schrader, and C.
scabriuscula (Pino-Bodas et al. 2015). No new sequences generated.
Specimen examined & observations. – CANADA. ALBERTA: Edmonton, 53.51078, -113.46508,
2019, on soil, T.L. Dueck (unvouchered observation: NatureLynx record https://
naturelynx.ca/sightings/14146/details); Edmonton, near Northland sandpit, 2 mi W and 1 mi S of 170 St.
and 79 Ave., 1977, on soil, D.C. Lindsay s.n. (PMAE-B77.24.107); Edmonton, River Loop Trail south of
Fort Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland 2021-20 & S. Toni (hb.
Haughland).
Cladonia pyxidata (L.) Hoffm. FIGURE 52 E-F.
River valley terricole and epiphyte. Characterized by short (to 3 cm), broadly cupped podetia
arising from persistent, appressed, well-developed primary squamules. The podetia are esorediate, but the
cups can be granular or patchy-corticate on the outside. A diagnostic feature is the peltate, pancake-like
squamules found lining the inside of the cups. It can be difficult to distinguish from mature Cladonia
chlorophaea especially as they grow intermingled on a variety of substrates within Edmonton and across
Alberta—see the entry for that species for information on some similar species. Additional PD+ orange
species to consider include C. pocillum (Ach.) O. J. Rich. and C. magyarica Vain., which are more
common on soil in arid parts of the province. Cladonia pocillum has relatively thick primary squamules
with a chalky, thick medulla. Cladonia magyarica tends to be more grey-green with less brown coloration,
often has secondary squamules arising from the cup margins, and it is K+ yellow due to the presence of
atranorin. Chemistry: K- or K+ dingy brown, KC-, C-, UV254-, PD+ orange. Secondary metabolites
detected by TLC: fumarprotocetraric acid. Molecular support: unresolved, sequences mapping to multiple
branches throughout the Clade “Cladonia”, subclade “Graciles” (Stenroos et al. 2018). No new sequences
generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Hawrelak Trail off-
leash park, 53.520733, -113.54158, 2020, on Betula base, D. Haughland 2020-19B (hb. Haughland);
Edmonton, Patricia Ravine, 53.503638, -113.593736, 2020, on soil along roots of live, large Picea glauca,
D. Haughland & A. Hood (unvouchered observation); Edmonton, near Northland sandpit, 2 mi W and 1 mi
S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.191).
Cladonia scabriuscula (Delise) Nyl. FIGURE 53 D.
Ravine system terricole and xylicole. Primary thallus not observed, forming distinctive podetia
that are 3–9 cm tall, typically pale at base, green to whitish at top, mostly slender, with open axils and
branching tips. The podetia are smoothly corticate except in the upper half to third where they become
scabrose and patchily granular-sorediate. Chemistry: K- or K+ dingy brown, KC-, C-, UV254-, PD+ orange.
Secondary metabolites detected by TLC: fumarprotocetraric acid. Molecular support: unresolved and
polyphyletic, in a complex with C. farinacea, C. furcata, and C. multiformis (Pino-Bodas et al. 2015,
Stenroos et al. 2018). No new sequences generated.
Observation. – CANADA. ALBERTA: Edmonton, Larch Sanctuary, 53.447302, -113.550894,
2020, terricolous, D. Haughland (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/10685/details); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315,
2021, on decayed stump, D. Haughland 2021-23A (hb. Haughland).
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Cladonia subulata (L.) F. H. Wigg. FIGURE 53 E.
Apparently rare ravine system terricole/xylicole. Primary squamules small to evanescent. Wand-
like podetia grey-green to brownish-green, sometimes paler at base, to 5 cm tall locally, typically entirely
sorediate. The podetia may be uniform in width and terminate in a blunt tip, form narrow cups, often with
marginal proliferations, or branch in bifurcations that can resemble antlers. See Cladonia. conocraea entry
for tips to discriminate similar species. Chemistry: K- or K+ dingy brown, KC-, C-, UV254-, PD+ orange.
Secondary metabolites detected by TLC: fumarprotocetraric acid. Molecular support: monophyletic with
strong support based on 3–15 sequences (Pino-Bodas et al. 2010, Stenroos et al. 2018).
Observation. – CANADA. ALBERTA: Edmonton, MacKenzie Ravine, by boardwalk, 53.52914,
-113.5603, 2020, on Picea glauca snag, D. Haughland 2020-110 (hb. Haughland); Edmonton, River Loop
Trail S of Fort Edmonton, 53.500627, -113.576611, 2021, on trailside soil, D. Haughland 2021-15 & S.
Toni (hb. Haughland).
GROUP 15. EPIPHYTIC FRUTICOSE LICHENS
Ten species. Key literature: Bird 1974; Brodo et al. 2001; Brodo 2016; Brodo & Hawksworth
1977; Clerc 2011; Halonen et al. 1998; Mark et al. 2016b; Wylie 1977. For Bryoria, we retain the
ecological and morphological species concepts of Brodo and Hawksworth (1977) versus recent
phylogenetic species concepts (Boluda et al. 2019, Velmala et al. 2014). Similarly, we use the
morphospecies concepts of Usnea from Clerc (2011) for the few species documented in Edmonton rather
than treat isidiate specimens historically called U. substerilis as synonymous with U. lapponica (as per
Mark et al. 2016b, now called U. perplexans, Clerc 2016). We do not doubt the veracity of recent
molecular work or nomenclatural corrections; rather we suspect that adopting taxonomic shifts here and in
related work across the province would be more confusing than illuminating at this time, especially given
the lack of resolution in studies to date. We provide limited spot test results as we seldom use them to help
discriminate Usnea species; we have found them to be uninformative due to overlapping chemotypes or
unreliable because of variable concentrations of metabolites. An additional epiphytic fruticose lichen
reported which we could not confirm despite numerous attempts in the field and in the herbarium is
Pseudevernia consocians (Whitemud Creek near ski slope, Edmonton, 1963, A.W. Stewart 575 [ALTA]),
Government of Alberta 2020). This species is more common in eastern North America but it has
confirmed, disjunct populations as far west as Alberta (ABMI 2020).
1a. Thallus hair-like, brown ............................................................................................... Bryoria fuscescens
1b. Thallus hair-like to shrub-like, greenish yellow ........................................................................................ 2
2a. Thallus with a tough, elastic, cartilaginous central cord (stretch gently along axis to see cord) ... 3 Usnea
2b. Thallus lacking a cord ............................................................................................................................... 6
3a. Thallus lacking isidia, soralia present, eroding the branch when fully developed, with flaps of cortex
around the soralia ................................................................................................................ Usnea perplexans
3b. Thallus isidiate, either individually from the branch in esorediate species or arising from larger soralia
mixed with soredia (may be sparse so check soralia carefully) ....................................................................... 4
4a. Thallus elongate; branches growing relatively long and parallel to each other ................. Usnea scabrata
4b. Thallus shrubby; branches diverging from each other at wide angles ....................................................... 5
5a. Soralia variable but wider than half the branch width, developing sparse to abundant isidia within the
soralia; attachment point black; papillae present at base ................................................. Usnea substerilis s.l.
5b. Well-defined soralia lacking, instead developing punctiform isidia over the surface; attachment point
typically pale; lacking papillae ...................................................................................................... Usnea hirta
6a. Thallus soft and pliable (cortex thin), dull, typically wrinkled; coarse soredia and/or isidia arising along
the branches; medulla cottony ....................................................................................... Evernia mesomorpha
6b. Thallus stiff (cortex thick), often shiny, smooth; if soralia present, in well-defined soralia rather than
occurring throughtout the thallus; medulla dense to honey-comb-like ........................................... 7 Ramalina
145
7a. Soralia lacking, apothecia typically present .............................................................................................. 8
7b. Soralia present along lobe edges or on lobe tips; apothecia not observed ................................................. 9
8a. Thallus small (1–2 cm long), inflated and spiny, partially hollow and perforate, somewhat translucent
......................................................................................................................................... Ramalina dilacerata
8b. Thallus larger (to 3 cm long), flattened and solid, branches fan-shaped, imperforate ... Ramalina sinensis
9a. Thallus inflated at least in part, with perforations into hollow branch interior; soredia present within
hooded soralia or on tattered lobe tips ................................................................................ Ramalina obtusata
9b. Thallus solid, flattened; soralia variable but mostly discrete, marginal and terminal, becoming laminal
on older tips ..................................................................................................................... Ramalina pollinaria
Bryoria fuscescens (Gyelnik) Brodo & D. Hawksw.
(= Bryoria lanestris (Ach.) Brodo & D. Hawksw.) FIGURE 55 A.
River valley and ravine system epiphyte. Growing to 5–8 cm in length in Edmonton, Bryoria
fuscescens is greyish-brown to dark-brown, and typically has abundant pale-green, fusiform soralia that are
PD+ orange. An abundant epiphyte particularly on conifers in northern Alberta, this species is sparse
locally. This is the only Bryoria documented to date in Edmonton. Chemistry: soralia PD+ orange, thallus
PD-, all other spot tests negative. Secondary metabolites detected by TLC: fumarprotocetraric acid.
Molecular support: high species-level support with recent reduction of two species to synonymy with B.
fuscescens (Boluda et al. 2019): B. vrangiana (Gyelnik) Brodo & D. Hawksw. (morphologically
distinguished by pseudocyphellae and more robust, regular branches), and B. capillaris (Ach.) Brodo &
Hawksw. (distinguished by lack of soralia in North America and presence of alectorialic acid and
barbatolic acid). Synonymy not adopted here. No new sequences generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Patricia Ravine,
53.503216, -113.592645, 2020, on live Prunus stems, D. Haughland 2020-44 & A. Hood (hb. Haughland);
Edmonton, Patricia Ravine, 53.502112, -113.592209, 2020, epiphytic, S. Toni (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/12139/details); Edmonton, Rio Park, 53.50320938, -
113.592617, 2020, epiphytic, L. Hjartarson (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/12116/details); Edmonton, MacKenzie Ravine, 1976, D.C. Lindsay s.n
(PMAE-B77.24.174 ); Edmonton, near Northland sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977,
on rotten wood, D.C. Lindsay s.n. (PMAE-B77.24.92); Edmonton, grassy park next to Saskatchewan Drive
bordering river valley, 53.513197, -113.53866, 2021, on Picea twigs, D. Haughland 2021-33C (hb.
Haughland).
Evernia mesomorpha Nyl. FIGURE 55 B.
Occasional river valley and ravine system and parkland epiphyte. One of the most common
epiphytes in forested regions of Alberta, this species is rare in Edmonton outside of river valley parks.
Thalli forming pale-green shrubby tufts of wrinkled branches bearing soredia and isidia along the ridges.
The dull, wrinkled outer cortex and cottony medulla help separate this genus from Usnea (compact and
often shiny cortex and elastic, cartilaginous central cord) and Ramalina (shiny cortex, often perforate or
with linear pseudocyphellae, and a solid to honey-combed interior). No other Evernia species are found in
Alberta’s Boreal or Parkland Natural Regions (ABMI, unpub.. Chemistry: cortex KC+ yellow, medulla
UV+ white, all other spot tests negative. Secondary metabolites detected by TLC: usnic acid (not detected
at low concentrations), divaricatic acid,. Molecular support: The genus originally appeared polyphyletic
(Crespo et al. 2010) or paraphyletic (Piercey-Normore 2006). In recent analyses, three species of Evernia
formed a monophyletic clade (Divakar et al. 2017). More work is needed to ascertain whether the species is
monophyletic as E. mesomorpha forms a clade with E. esorediosa (Müll. Arg.) Du Rietz in some analyses
(Piercey-Normore 2006). No new sequences generated.
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Urban
Monitoring Site 4, 53.443571, -113.461517, 2019, on trunk of Ulmus americana, J. Birch & J. Wasyliw
[UoA-CC-72] (hb. Haughland); Edmonton, Patricia Ravine, 53.503216, -113.592645, 2020, on Prunus
stems, D. Haughland 2020-47 & A. Hood (hb. Haughland); Whitemud Park, Edmonton, 53.497703, -
113.561303, 2020, epiphytic, hanna1025 (unvouchered observation: iNaturalist record https://www.inatur-
146
Figure 55. Epiphytic fruticose lichens of Edmonton plate 1. A, Bryoria fuscescens, Toni
https://naturelynx.ca/sightings/12139/details. B, Evernia mesomorpha, Edmonton, Buena Vista, 2020,
Haughland unvouchered observation. C, Ramalina dilacerata, wet thallus, Haughland 2020-3. D,
Ramalina obtusata, wet thallus, Haughland 2020-5. E, Ramalina pollinaria, wet thallus, Haughland 2020-
14. F, Ramalina sinensis, Haughland https://naturelynx.ca/sightings/14278/details.
147
-alist.org/observations/59078955); Edmonton, between Stony Plain Rd. and 100 Ave. at 148 St., 1976, on
supports of wooden footbridge, D.C. Lindsay s.n. (PMAE-B77.24.141); Edmonton, MacKenzie Ravine,
1976, on wood, D.C. Lindsay s.n. (PMAE-B77.24.37); Edmonton, near Northland sandpit, 2 mi W and 1
mi S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.99).
Ramalina dilacerata (Hoffm.) Hoffm. FIGURE 55 C.
River valley and ravine system epiphyte. A commonly overlooked but frequent boreal species
characterized by small (1–2 cm) shrubby, tufted thalli. The cortex is shiny and semi-translucent and lacks
the pseudocyphellae present in other species of the genus (e.g., Ramalina sinensis). Upon close
examination, it is easily identified by its hollow, slightly inflated branches that develop characteristic
punctures on the underside. Apothecia abundant, relatively large, with pale pinkish-yellow, lightly pruinose
discs, forming near the ends of branches. No vegetative propagules. Chemistry: cortex KC+ yellow,
medulla UV+ whitish, all other spot tests negative. Secondary metabolites detected by TLC: usnic acid (not
detected at low concentrations), divaricatic acid. Molecular support: monophyletic with high support in an
analysis of five sequences (Timsina et al. 2012). No new sequences generated.
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek,
53.502937, -113.602263, 2020, stems of Corylus cornuta, D. Haughland 2020-3 & C. Shier (hb.
Haughland); Edmonton, Patricia Ravine, 53.502112, -113.592209, 2020, epiphytic, S. Toni (unvouchered
observation: NatureLynx record https://naturelynx.ca/sightings/12144/details); Edmonton, MacKenzie
Ravine, 1976, on rotting log, D.C. Lindsay s.n. (PMAE-as minor component in B77.24.41).
Ramalina obtusata (Arnold) Bitter FIGURE 55 D.
Apparently rare river valley and ravine system epiphyte. A boreal species often found growing
with Ramalina pollinaria elsewhere in Alberta. Characterized by yellowish-green, shrubby, shiny thalli,
growing in tufts to 3 cm long. The branches usually are flattened near the attachment point, and become
hollow and slightly inflated with open branch tips that terminate in tattered to hooded soralia. Distinguished
from R. pollinaria by its inflated branches (vs. solid and flattened in R. pollinaria) and the terminal soralia
(vs. marginal and terminal in R. pollinaria). Chemistry: cortex KC+ yellow, all other spot tests negative.
Secondary metabolites detected by TLC: usnic acid (not detected at low concentrations), evernic acid
and/or obtusatic acid (cannot differentiate solvents used herein). Molecular support: lacking, too few
sequences available at present. Polyphyletic within Ramalina in a recent analysis including two sequences
(Haughland, in prep.). No new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Wolf Willow Creek, 53.502787, -
113.601791, 2020, on bark of dead Picea glauca, D. Haughland 2020-5 & C. Shier (hb. Haughland);
Edmonton, Wolf Willow Creek, 53.502228, -113.60128, 2020, on Salix, D. Haughland 2020-13A & C.
Shier (hb. Haughland); Edmonton, MacKenzie Ravine, 1976, on bark of twig, D.C. Lindsay s.n. (PMAE-
B77.24.173); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315, 2021, on Betula papyrifera, D.
Haughland 2021-22C (hb. Haughland).
Ramalina pollinaria (Westr.) Ach. FIGURE 55 E.
Occasional river valley and ravine system epiphyte. A common and variable boreal species.
Thallus yellowish-green, shrubby, dull to slightly shiny, growing in tufts to 3 cm long. Branches usually
flattened, solid throughout, slightly longitudinally striate. Soredia farinose, in delimited soralia along the
lobe margins and in terminal, slightly labriform soralia. Apothecia not found. See Ramalina obtusata entry
for points of distinction. Gasparyan et al. (2017) described R. labiosorediata Gasparyan, Sipman &
Lücking as a segregate from R. pollinaria in North America. While R. labiosorediata has not been detected
in Alberta, it is distinguished by relatively broad lobes that have few to no marginal soralia and well-
developed terminal labriform soralia. Chemistry: cortex KC+ yellow, all other spot tests negative.
Secondary metabolites detected by TLC: usnic acid (not detected at low concentrations), evernic acid
and/or obtusatic acid (cannot differentiate in solvents used herein). Molecular support: high at species level.
Recent analyses show that a sequence from Edmonton (isolate DLH12 from Haughland 2020-13B) and two
collections from northern Canada form a highly supported, monophyletic clade with R. pollinaria s.s. from
148
Gasparyan et al. (2017), not with R. labiosorediata, evidence that both species occur in North America
(Haughland et al. in prep.).
Representative specimens examined & observations. – CANADA. ALBERTA: Edmonton, Wolf
Willow Creek, 53.502228, -113.60128, 2020, on Salix, D. Haughland 2020-13B & C. Shier (hb.
Haughland); Edmonton, Whitemud Ravine, 53.495491, -113.55983, 2020, on deciduous tree bark, D.
Haughland 2020-14 & P. Williams (hb. Haughland); Edmonton, Patricia Ravine, 53.503105, -113.592863,
2020, on bark of downed, intact Betula papyrifera, D. Haughland 2020-41 & A. Hood (hb. Haughland);
Edmonton, Kinnaird Ravine, 53.558953, -113.459253, 2020, on Picea glauca snag, D. Haughland 2020-30
& P. Williams (hb. Haughland); Edmonton, Patricia Ravine, 53.503448, -113.593252, 2020, epiphytic, L.
Hjartarson (unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/12095/details);
Edmonton, Patricia Ravine, 53.502112, -113.592209, 2020, epiphytic, S. Toni (unvouchered observation:
NatureLynx record https://naturelynx.ca/sightings/12450/details); Edmonton, Whitemud Creek Ravine,
53.495491, -113.559830, 2020, epiphytic, D. Haughland (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/12606/details); Edmonton, Whitemud Creek Ravine, 53.494633, -
113.560377, 2020, epiphytic, D. Haughland (unvouchered observation: NatureLynx record
https://naturelynx.ca/sightings/12607/details); Edmonton, MacKenzie Ravine, 1976, on tree, D.C. Lindsay
s.n. (PMAE-B77.24.26); Edmonton, Terwillegar Footbridge, 53.4797, -113.594315, 2021, on Betula
papyrifera, D. Haughland 2021-22B (hb. Haughland); Edmonton, MacKenzie Ravine, by boardwalk,
53.52914, -113.5603, 2020, on Picea glauca snag, D. Haughland 2020-109 (hb. Haughland).
Ramalina sinensis Jatta FIGURE 55 F.
Rare tablelands epiphyte. This species is characterized by cartilaginous flattened lobes that tend to
be deeply dissected, and at maturity terminate in large pinkish-yellow apothecia. The lobes attach at a
single holdfast, and have long pseudocyphellae on one surface, developing between the cartilaginous ribs
that run parallel to the branch in well-developed specimens. The single specimen collected in Edmonton
resembled Ramalina unifolia J.W. Thomson (distinguished by a lack of dissection, forming thalli composed
of a single, fan-like lobe that otherwise resemble R. sinensis (Thomson 1990)). Because we are uncertain
about the validity of that species, we retain this sample within R. sinensis. Molecular support: mixed. An
early-diverging species within Ramalina with at least three highly supported lineages (one from Alberta)
within a deeper, monophyletic species-level clade; some authors suggest more research is needed to
determine whether these branches should be split into separate species (Spjut et al. 2020, Timsina et al.
2012). In contrast, LaGreca et al. (2020) found that R. sinensis is paraphyletic with a strongly supported
clade including two sequences of R. unifolia. More work is needed to determine if R. unifolia should be
reduced to synonymy with R. sinensis. No new sequences generated.
Specimen examined. – CANADA. ALBERTA: Edmonton, Urban Monitoring Site 9, 53.453708, -
113.527608, 2019, on trunk of Fraxinus, D. Haughland & A. Stordock s.n. [UoA-CC-76] (hb. Haughland);
Edmonton, Urban Monitoring Site 77E, 53.501971, -113.348115, 2019, on trunk of Populus, 2019, D.
Thauvette & J. Birch (unvouchered observation).
Usnea hirta (L.) Weber ex F. H. Wigg. FIGURE 56 A.
River valley epiphyte. A species characteristic of northern coniferous forests in Alberta, this
shrubby species typically is densely branched, forming compact tufts up to 5 cm long. The branches are
ridged and/or foveolate, lack papillae or soredia, and have abundant punctiform isidia (arising singly from
the branch). Some specimens have abundant fibrils, which resemble small, perpendicular side branches.
The attachment point typically is pale, whereas the outer-most branch tips often are blackened. May be
confused with Usnea. scabrata, but that species has a blackened attachment point and typically abundant,
obvous papillae, especially near the base. Chemistry: PD-, K-, KC+ yellow, C-, UV-. Secondary
metabolites detected by TLC: usnic acid. Molecular support: limited to date. Two sequences (both from
Scotland) are monophyletic with strong support (Truong et al. 2013). No new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Hawrelak Trail off-leash park,
53.519809, -113.540503, 2020, on Betula bark, D. Haughland 2020-17 (hb. Haughland); Edmonton,
Patricia Ravine, 53.503216, -113.592645, 2020, on Prunus stems, D. Haughland 2020-46 & A. Hood (hb.
Haughland); Edmonton, near Northland sandpit, 2 mi W and 1 mi S of 170 St. and 79 Ave., 1977, on wood,
D.C. Lindsay s.n. (PMAE-as minor component in B77.24.96).
149
Figure 56. Epiphytic fruticose lichens of Edmonton plate 2: Usnea. A, U. hirta, Redwater Natural Area
northeast of Edmonton, Haughland https://naturelynx.ca/sightings/15781/details. B, U. perplexans, closeup
of excavate soralia, ca. 12 km NW of Zama City, ABMI Site 158, 59.23, -118.86, 2014, epiphytic, D.
Hogarth s.n. [ABMI Lichen # 577649]. C, U. substerilis, Haughland 2020-94. D, U. scabrata, Haughland
unvouchered observation.
Usnea perplexans Stirt.
(= Usnea lapponica Vain.) FIGURE 56 B.
River valley epiphyte. Characterized by short, shrubby thalli up to 8 cm long, but typically shorter
in the city. The branches can be round or dented and are densely papillate; the branches typically diverge
but may grow parallel to each other in more luxurious morphs outside of Edmonton. It is best characterized
by the development of deeply-excavate soralia that develop flaps of cortex around the soralia as the branch
erodes, as well as the lack of isidia. The attachment point is typically black. Similar shrubby species
include Usnea substerilis (with at least some isidia within the soralia, common) and U. glabrescens (Nyl.
ex Vain.) Vain. (with round, excavate soralia that contain sparse isidia only when young; a well-developed,
tree-like attachment point (resembling roots flaring from the base of a tree trunk and anchoring the thallus
to the substrate; thick cortex and thin medulla in longitudinal section; and typically with norstictic acid, rare
in Alberta). Chemistry: spot tests variable, not diagnostic. Secondary metabolites and chemotypes detected
by TLC: usnic acid only (63%), usnic acid with salazinic acid (31%), usnic acid with barbatic acid (4%), or
usnic acid with salazinic acid and barbatic acid (2%). These chemotypes agree with those documented
elsewhere (e.g., Halonen et al. 1998), but recent phylogenetic analyses suggest that specimens with barbatic
acid may belong to U. wasmuthii Räsänen (Mark et al. 2016b). Molecular support: weak at species-level.
Clades in a multi-locus phylogeny do not correspond to traditionally circumscribed species. Because of the
lack of resolution in a clade of U. lapponica, U. substerilis, U. barbata (L.) F.H. Wigg. and U. intermedia
150
(A. Massal.) Jatta, Mark et al. (2016b) placed U. substerilis in synonymy with this species. Synonymy not
adopted at this time; no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, Hawrelak Trail off-leash park,
53.520721, -113.54155, 2020, on Betula bole, D. Haughland 2020-22 (hb. Haughland); Edmonton, Patricia
Ravine, 53.503216, -113.592645, 2020, on Prunus stems, D. Haughland 2020-45 & A. Hood (hb.
Haughland); Edmonton, MacKenzie Ravine, 1976, on tree, D.C. Lindsay s.n. (PMAE-B77.24.25);
Edmonton, grassy park next to Saskatchewan Drive bordering river valley, 53.513197, -113.53866, 2021,
on Picea twigs, D. Haughland 2021-33B (hb. Haughland).
Usnea scabrata Nyl.
(? = Usnea barbata (L.) F. H. Wigg.) FIGURE 56 C.
River valley epiphyte. Alberta’s most common pendant Usnea. Characterized by relatively long
branches that soon grow parallel to each other, typically branched very close to the base, with a narrowed,
blackened attachment point and a relatively thin axis (axis: medulla ratio ≤1.5). Fibrils may be sparse or
common. The branches are typically abundantly papillate, foveolate and ridged, with punctiform soralia
giving rise to isidia. Usnea dasopoga differs in its relatively thick axis relative to the medulla, its broad
attachment with numerous “root-like” extensions clasping the wood, and lack of foveoles. Chemistry: PD-,
K-, KC+ yellow, C-, UV-. Secondary metabolites detected by TLC: usnic acid. An additional metabolite
documented in this species elsewhere is salazinic acid (e.g., Clerc 2011). Molecular support: weak at
species-level. Clades in a multi-locus phylogeny do not correspond to traditionally circumscribed species
(Mark et al. 2016b).
Specimens examined & observations. – CANADA. ALBERTA: Edmonton, Patricia Ravine,
53.503216, -113.592645, 2020, on Prunus stems, D. Haughland & A. Hood (unvouchered observation);
Edmonton, Mill Creek Ravine South, 53.508933, -113.461392, 2019, on deciduous trees, T.L. Dueck
(unvouchered observation: NatureLynx record https://naturelynx.ca/sightings/9832/details); Edmonton,
grassy park next to Saskatchewan Drive bordering river valley, 53.513197, -113.53866, 2021, on Picea
twigs, D. Haughland 2021-33A (hb. Haughland).
Usnea substerilis Motyka s.l. FIGURE 56 D.
River valley epiphyte. Another short, shrubby species similar to Usnea perplexans, U. substerilis
traditionally is differentiated by soralia that vary from excavate to tuberculate but which seldom develop
cortical flaps, and instead give rise to sparse to abundant isidia. Separated from the morphologically
similar, rarer U. subfloridana Stirt. by chemistry (the latter with medulla UV+ white due to presence of
squamatic acid in Alberta). Chemistry: spot tests variable, not diagnostic. Secondary metabolites and
chemotypes detected by TLC: usnic acid only (40%), usnic acid with salazinic acid (35%), usnic acid with
barbatic acid (9%), usnic acid with salazinic acid and barbatic acid (14%). These chemotypes agree with
those documented elsewhere (e.g., Halonen et al. 1998), but recent phylogenetic analyses suggest that
specimens with barbatic acid may belong to U. wasmuthii (Mark et al. 2016b). Molecular support: weak at
species-level. Clades in a multi-locus phylogeny do not correspond to traditionally circumscribed species.
Because of the lack of resolution in a clade of U. lapponica, U. substerilis, U. barbata, and U. intermedia,
Mark et al. (2016b) placed U. substerilis in synonymy with U. lapponica (now U. perplexans). Synonymy
not adopted at this time; no new sequences generated.
Specimens examined. – CANADA. ALBERTA: Edmonton, near Northland sandpit, 2 mi W and 1
mi S of 170 St. and 79 Ave., 1977, on wood, D.C. Lindsay s.n. (PMAE-B77.24.96); Edmonton, Buena
Vista Meadow, 53.513721, -113.54878, 2020, on Picea glauca branch, D. Haughland 2020-94 & P.
Williams (hb. Haughland).
ACKNOWLEDGEMENTS
We are grateful to the generous lichenologists that have examined Alberta material for the senior author over
the years, helping inform many of the species concepts used here, including: Trevor Goward (Bryoria, Cladonia,
Peltigera, Usnea), Steve Selva (calicioids), Martin Westberg (Candelaria), Irwin Brodo (Cladonia, Lecanora,
Physcia), Teuvo Ahti (Cladonia), Matthias Shultz (Collema, Lichinaceae), James Lendemer (Heterodermia), Ted
Esslinger (Melanelia s.l., Phaeophyscia, Physcia, Physconia), and the Duke Peltigera lab (François Lutzoni, Jolanta
Miadlikowska, Ian Medeiros, and Carlos Pardo-De la Hoz). Special thanks to Toby Spribille for helpful conversation
151
about the project, providing ITS sequences through his lab, and assistance with Alyxoria and Bilimbia. We are also
grateful to A. Deneka for sharing information from ongoing work on a modern lichen catalog for the prairie provinces.
Our thanks to Jordan Bell and the ABMI for assistance with NatureLynx observations. Finally, we are grateful to the
nature app contributors for their observations, to Tiffany L. Dueck for permitting us to use some of her images, to the
City of Edmonton for a permit to collect lichens, to the two anonymous reviewers for their insights, corrections, and
encouragement, and to the editors of Opuscula Philolichenum for improving the quality of the keys and texts
substantively.
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SUPPLEMENTARY APPENDICIES
Supplementary Appendix 1: Species List. – The complete species list, including species that we could
not confirm, are included; in addition many traits for the lesser-known crustose species have been tabulated
as well so the table can serve as an interactive key.
Deposited in Dryad: https://doi.org/10.5061/dryad.sqv9s4n6d
Supplementary Appendix 2: Sequence Voucher Table. – The complete list of 456 sequences used in this
study, including the analyses each sequence was used in, the source of the sequence, voucher information
including herbarium where those data were available, collection location, publication the sequence was first
generated for, and GenBank numbers by genetic marker.
Deposited in Dryad: https://doi.org/10.5061/dryad.sqv9s4n6d
Supplementary Appendix 3: Species Distribution Maps. – Distribution maps for the confirmed historic
and extant ocurrences of all the species within Edmonton and treated here are presented on the following
pages. The maps are derived from the same basemap used in Figure 1. As most Usnea specimens were too
poorly developed for species-level identification, open circles on the Usnea maps indicate genus-level
records.
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