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Lichens and allied fungi of Salmonier Nature Park,
Author(s): Richard Troy McMullin and Yolanda F. Wiersma
Source: The Journal of the Torrey Botanical Society, 144(3):357-369.
Published By: Torrey Botanical Society
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Journal of the Torrey Botanical Society 144(3): 357–369, 2017.
Lichens and allied fungi of Salmonier Nature Park, Newfoundland
Richard Troy McMullin
Research and Collections, Canadian Museum of Nature, Ottawa, ON, K1P 6P4, Canada,
Department of Biology, Memorial University, St. John’s, NL, A1B 3X9, Canada
Yolanda F. Wiersma
Department of Biology, Memorial University, St. John’s, NL, A1B 3X9, Canada
Abstract. We conducted the first comprehensive floristic study of the lichens and allied fungi of Salmonier Nature
Park on the Avalon Peninsula in Newfoundland, Canada. By comparing our results to those from other provincial
parks in Newfoundland, we show that Salmonier Nature Park has a regionally rich lichen biota that includes several
uncommon species. We carry out an assessment of landscape-level drivers including geographic location and land
cover diversity to determine whether lichen richness corresponds to patterns at the landscape extent. Within
Salmonier, one species (Erioderma pedicellatum) is listed as ‘‘special concern’’ by the federal Committee on the
Status of Endangered Wildlife in Canada and ‘‘critically endangered’’ by the International Union for Conservation of
Nature. Two species are new to the island of Newfoundland: Phaeophyscia ciliata and Stereocaulon subcoralloides.
Six species are new to the province of Newfoundland and Labrador: Ephebe hispidula, Muellerella lichenicola,
Mycoblastus sanguinarioides, Placynthium flabellosum, Usnea flammea, and Xanthoparmelia angustiphylla. Our
results provide baseline knowledge that allows changes in the lichen community to be monitored, the discovery of
new species in the park to be acknowledged, regional distributions and frequencies to be better understood, and
accurate comparisons to be made with other parks.
Key words: Atlantic Canada, Avalon Forest Ecoregion, biogeography, landscape ecology, lichen biodiversity
Floristics creates fundamental knowledge that
provides an important foundation for conservation
initiatives (Reid and Miller 1989; Environment
Canada 1995; Powell, Barborak, and Rodriguez
2000). This information shapes our understanding
of species distribution and frequency in particular
areas. Comprehensive ﬂoristic studies also provide
a baseline from which changes can be monitored,
including acknowledging previously unreported
species or the effects of disturbances such as air
pollution or acid rain (Henderson 2000, Richard-
son and Cameron 2004, McMullin and Ure 2008).
Accurate ecological comparisons with other re-
gions or localities are made possible as well, which
contributes greater insight into landscape-level
diversity patterns (Pickett 1989, Fukami and
Wardle 2005). Lichens are particularly sensitive
to microclimatic conditions and are strong indica-
tors of variation among ecosystems (Brodo, Sharn-
off, and Sharnoff 2001; Nash 2008). However,
fundamental baseline knowledge for lichens in
many areas of Canada is limited or does not exist
(Goward, Brodo, and Clayden 1998).
On the Canadian island of Newfoundland,
lichens collections have been frequent relative to
other regions in the country (Ahti 1974, 1983;
Goward, Brodo, and Clayden 1998). Collections
date back to those made by Joseph Banks in 1766
(Lysaght 1971). Major published collections that
followed include those by Arthur Waghorne
(Arnold 1896, Eckfeldt 1895, Macoun 1902,
Brassard 1980), Teuvo Ahti (Ahti 1983), and John
McCarthy (McCarthy, Driscoll, and Clayden
2015). Important collections have also been made
that were not published (e.g., see Goward, Brodo,
and Clayden 1998; Pitcher and Clayden 2007).
Summaries of all species reported or collected in
Newfoundland were compiled by Eckfeld (1895)
and Ahti (1983), and an updated list is currently
We gratefully acknowledge D. Howell, J. Kennedy,
M. Blackwood, R. Jarvis, and the staff at Salmonier
Nature Park for hosting R.T.M. for the duration of this
study and providing permission for us to collect; K.
Unger and M. Lafferty, our ﬁeld companions from the
Nature Conservancy of Canada; J. Kennedy for being a
guide through unmarked wilderness; J. McCarthy for
assisting with determining new provincial and
Newfoundland records; M. Pitcher for helpful
discussion about the ecosystems and lichen species in
the park; B. Dorin for data entry; and R. Wigle for
curating specimens. This study was conducted under
provincial park research permit 0895. Funding was
provided by the Natural Sciences and Engineering
Research Council of Canada to Y.F.W.
Author for correspondence: tmcmullin@mus-nature.
ÓCopyright 2017 by The Torrey Botanical Society
Received for publication July 29, 2016, and in revised
form November 6, 2016; ﬁrst published June 30, 2017.
being assembled and added to by John McCarthy,
Teuvo Ahti, and Stephen Clayden. A history of
lichen collecting on the island is described by Ahti
(1974, 1983) and by Goward, Brodo, and Clayden
(1998). Since 1998, several additional lichen
collections from Newfoundland have been pub-
lished (Deduke and Piercey-Normore 2013; Pier-
cey-Normore 2013; Deduke, Ahti, and Piercey-
Normore 2014; McCarthy, Driscoll, and Clayden
2015; McMullin and Arsenault 2016). In 2007, an
international group of lichenologists gathered for
the Tuckerman Workshop and made collections at
several sites on the Avalon Peninsula (the Avalon)
(Pitcher and Clayden 2007), which comprises the
easternmost region of Newfoundland and is known
for a rich and unusual lichen biota (Ahti 1974,
1983; McCarthy, Driscoll, and Clayden 2015;
COSEWIC 2014). Despite the large number of
lichen collections that have been made in New-
foundland, comprehensive inventories of spatially
deﬁned areas (e.g., parks and protected areas) have
been limited and new island records are still
regularly discovered (Knudsen and Kokourkova
2015; McCarthy, Driscoll, and Clayden 2015;
McMullin and Arsenault 2016).
A recent survey of lichen diversity in New-
foundland (McCarthy, Driscoll, and Clayden
2015) found 133 taxa, of which 15 were new
records for the province. McCarthy, Driscoll, and
Clayden (2015) surveyed four provincial parks,
which extended from the west to the east coast of
the island, in close proximity to the southern coast
of the island. Of these, one (Jipujijikeui Kues-
pem) is a known lichen hotspot, and Fitzgerald’s
Pond on the Avalon protects a population of the
globally rare lichen Erioderma pedicellatum
(Scheidegger 2003, COSEWIC 2014). Salmonier
Nature Park (Salmonier), located in the center of
the Avalon, is currently the only protected area in
the Avalon Forest Ecoregion, the smallest ecor-
egion in the province (Damman 1983), but which
is known for an unusual and rich lichen biota
(Ahti 1983). None of the provincial parks
surveyed by McCarthy, Driscoll, and Clayden
(2015) are in this ecoregion, but Salmonier Nature
Park is close to one of them (ca. 35 km east of
Fitzgerald’s Pond Provincial Park Reserve). To
our knowledge, a detailed list of the lichens of
Salmonier Nature Park has not been published.
Given the proximity of Salmonier to known areas
of high lichen richness (Ahti 1983), and its status
as a protected area that is largely unimpacted (see
Study Area below), it is valuable to gain a better
understanding of the lichen diversity within this
In addition to developing a comprehensive list
of lichen and allied fungi in Salmonier Nature
Park, we were interested in understanding patterns
of lichen diversity at larger spatial extents. A
comparison of lichen richness and species compo-
sition in the four parks surveyed by McCarthy,
Driscoll, and Clayden (2015) to our survey here
might yield insights into the alpha, beta, and
gamma diversity along a roughly east-west transect
of protected areas on the island of Newfoundland.
We hypothesize that factors that might affect
within-park species richness include park size
(driven by classic species-area relationships) and
habitat diversity within parks (parks with a higher
variety of land cover/habitat types will have higher
lichen species richness). We expect to see variation
in species composition as we move east to west,
driven largely by climatic and ecoregion differ-
Methods. STUDY AREA. Salmonier Nature Park
is a provincial protected area managed by the
Wildlife Division for the purpose of environmental
education (Fig. 1). The park was established in
1978 and is 1,455 ha in size. Of this, most visitors
only visit the 40 ha that include a boardwalk and
animal enclosures where the park houses injured or
orphaned wildlife. These are available for viewing
to the public for the purpose of increasing
understanding and appreciation of the province’s
wildlife (Government of Newfoundland and Lab-
rador, no date). The park receives over 40,000
visitors per annum; however, very few of these
visit the 1,415 ha of ‘‘backcountry’’ that exists
outside of the nature trail and educational facilities.
Other than two primitive cabins and a few trails
(both of which are primarily used by park staff),
the majority of the park is not heavily impacted by
visitor use. The eastern boundary of the nature
park borders on the 1,070-km
Reserve, the largest protected area on the Avalon
Peninsula, which represents a portion of the
Maritime Barrens Ecoregion.
Salmonier Nature Park is located within the
Avalon Forest Ecoregion, which is the smallest
ecoregion in the province (555 km
1983). This Ecoregion is characterized by ribbed
moraines and domed bogs of glacial origin,
covered with forests dominated by balsam ﬁr
358 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 144
FIG. 1. (A) Salmonier Nature Park. Polygons indicate habitat types (based on Earth Observation for
Sustainable Development data course for land cover for which a representative sample was surveyed in this
study). Roman numerals indicate sampling locations as described in Table 1. Colored polygons (see map legend
for description) indicate habitat types for which no representative site was sampled in this study. (B) Indicates
location of Salmonier Nature Park (diamond) relative to the four provincial parks (triangles) recently surveyed for
lichens. (C) Shows location of the province of Newfoundland and Labrador (dark grey) within Canada.
2017] MCMULLIN AND WIERSMA: LICHENS OF SALMONIER NATURE PARK 359
(Abies balsamea (L.) Mill.) intermixed with yellow
birch (Betula alleghaniensis Britt.) and white birch
(Betula papyrifera Marsh.). The climate is charac-
terized by cool summers, with a mean July
temperature of 16.5 61.5 8C; mild winters, with
a mean January temperature of 3.1 61.8 8C
(Environment Canada 2015); and high precipita-
tion, over 1,100 mm per year (Damman 1983).
SAMPLING. Our lichen survey at Salmonier
occurred between October 5 and October 29,
2015. Sampling was scattered throughout this
time, totaling approximately 4 days. Our survey
methods followed Newmaster et al. (2005), who
showed that examining large areas (referred to as
ﬂoristic habitat sampling) captures cryptogam
diversity more effectively than using smaller
representative plots. Using ﬂoristic habitat sam-
pling, we surveyed as many ecosystems in the park
as possible. All observed restricted mesohabitats
(e.g., streams, rock outcrops) were examined in
each ecosystem. We attempted to assess as many
microhabitats (e.g., snags, different tree species
and rock types) as possible at each location. This
method was described as an ‘‘intelligent meander’’
by Selva (1999, 2003).
IDENTIFICATION. We identiﬁed specimens using
microscopy to examine morphology and chemical
spot tests following Brodo, Sharnoff, and Sharnoff
(2001). Chemistry was further examined using an
ultraviolet light chamber and thin-layer chroma-
tography following Culberson and Kristinsson
(1970) and Orange, James, and White (2001) in
solvents A and C. Images were captured using a
Canon PowerShot Elph 130 IS digital camera.
Voucher specimens have been deposited at the
Ayre Herbarium at Memorial University and the
Canadian Museum of Nature.
GEOGRAPHIC INFORMATION SYSTEM ANALYSIS.To
contrast habitat diversity in Salmonier Nature Park
with the four provincial parks surveyed by
McCarthy, Driscoll, and Clayden (2015) we
quantiﬁed the land cover diversity using remotely
sensed satellite data. While the dataset used, the
Earth Observation for Sustainable Development
(EOSD) data (Wulder and Nelson 2003), does not
yield land cover classes that match the habitats
described by McCarthy, Driscoll, and Clayden
(2015) they provide an index of overall diversity of
land cover types. The EOSD data are based on
Landsat TM imagery and classiﬁed to the same
standard across the country with a 30-m pixel size.
We overlaid these data for the province with the
boundary ﬁles for the four provincial parks and for
Salmonier Nature Park (obtained from CanVec
geogratis/11042). We clipped the EOSD raster by
the boundaries of each park and then used the data
in each attribute table to count land cover types
(number of EOSD classes, not including shadow
and cloud), which we took to represent ‘‘land cover
richness’’ within each park. We also used the count
of pixels for each land cover type to calculate
landscape-level Shannon diversity index for each
park as follows:
is the proportion of pixels within the park
of land cover type i. We chose the Shannon index
over the Simpson’s index because it has been
shown in landscape analyses that the Shannon
index better captures and quantiﬁes diversity in
landscapes with rare habitat types (Nagendra
2002). Given the habitat speciﬁcity of many
lichens, we hypothesize that those parks with
more rare habitats or higher overall land cover
diversity will have higher lichen diversity.
SORENSEN-DICE COEFFICIENT OF SIMILARITY.We
used the Sorensen-Dice coefﬁcient of similarity to
compare the lichen community across all pair-wise
comparisons of the ﬁve parks. The coefﬁcient is
calculated as follows:
where Arepresents the total number of species
common to both parks, Bis the number of species
at park 1 that are absent from the other park, and C
is the number of species at park 2 that are absent
from park 1.
Results. LANDSCAPE DIVERSITY. Land cover
richness was highest in Salmonier and Fitzgerald’s
Pond parks, with 14 land cover classes in each.
Shannon landscape diversity was highest in
Sandbanks Provincial Park and lowest in Jipujij-
kuei Kuespem (Table 2).
SPECIES DIVERSITY. One hundred and thirty-seven
lichen and allied fungus species in 66 genera were
discovered at Salmonier during our study. Com-
bined with previous unpublished reports, which are
the nonbolded species in the annotated list below,
144 species in 67 genera are now known from the
360 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 144
park. These include 46 (32%) microlichens
(crustose species, which includes all allied fungi)
and 98 (68%) macrolichens (44 foliose and 54
fruticose). One hundred and twenty-seven (88%)
species have green algae as a primary photobiont,
12 (8%) species have cyanobacteria as their
primary photobiont, and 5 (3%) species are
nonlichenized fungi traditionally treated with
lichens. Seven (5%) species are calicioids and
two (1.4 %) species are lichenicolous.
The species with cyanobacteria as a primary
photobiont (cyanolichens) are Ephebe hispidula,
Erioderma pedicellatum (Fig. 2), Fuscopannaria
ahlneri, Lichinodium sirosiphoideum, Parmeliella
parvula, Peltigera canina, Peltigera hymenina,
Peltigera membranacea, Peltigera polydactylon,
Table 1. Lichen collection sites in Salmonier Nature Park. Roman numerals correspond with collection
numbers in the Annotated Species List.
Site Name Latitude Longitude Habitat
I Boardwalk Trail 47.263798N
Mature and humid mixed-wood
conifer forests. Tree cover
dominated by Abies balsamea
and Picea spp.
II Trail between the
and the old visitor center
47.263468N 53.285158W A mature mixed-wood conifer
forest dominated by Abies
balsamea and Picea spp.
surrounding a bog.
III Butler’s Pool 47.245848N 53.270838W A humid sheltered mixed-wood
conifer forest along the
Salmonier River. Tree cover
dominated by Abies balsamea
and Picea spp. Exposed
boulders are along the shores of
IV Boardwalk Trail, around
the snowy owl enclosure
47.26278N 53.283288W Sparsely treed bog. Tree cover
dominated by Picea mariana
and Larix laricina.
V Three Rivers 47.252838N 53.251098W Exposed boulders and shoreline
trees (conifers) among the three
rivers entering Three Rivers
Pond from Salmon Pond.
VI Mackay’s Lookout 47.245818N 53.237328W Maritime barrens with the highest
points of exposed rock reaching
VII Trail between the three
rivers and Metcalfe Falls
Mature mixed-wood forest
dominated by Abies balsamea,
Betula papyrifera, and Picea
VIII Murphy’s Falls 47.247288N 53.259668W Exposed boulders in the river
below a waterfall.
IX Trail from Highway 90 to
Mature mixed-wood forest
dominated Abies balsamea,
Betula alleghaniensis, and
X Whiskey Ponds River 47.251188N 53.286978W Exposed boulders in the river
between the two smallest ponds
in the Whiskey Ponds chain.
XI Eastern moraines 47.249818N 53.281348W Moraines in the eastern corner of
Salmonier Nature Park that are
surrounded by sparsely treed
bogs. Tree cover on the
moraines is dominated by Abies
balsamea and Picea mariana.
In the bogs, scattered tree cover
is dominated by Larix laricina.
XII Metcalfe’s Falls 47.2525378N 53.2329488W Exposed boulders in the river
above and below a waterfall.
2017] MCMULLIN AND WIERSMA: LICHENS OF SALMONIER NATURE PARK 361
Peltigera ponojensis,Peltigera scabrosa, and
Placynthium ﬂabellosum; the calicioid species are
Calicium abietinum, Calicium lenticulare, Chae-
notheca balsamconensis, Chaenotheca brunneola,
Mycocalicium subtile, Phaeocalicium compressu-
lum, and Phaeocalicium matthewsianum; and the
lichenicolous species are Cystobasidium hypogym-
niicola and Muellerella lichenicola.
SORENSEN-DICE COEFFICIENT OF SIMILARITY. Fitz-
gerald’s Pond (closest park to Salmonier) had the
highest similarity coefﬁcient relative to Salmonier
(Table 3). Sandbanks and J.T. Cheeseman are both
in the Maritime Barrens Ecoregion, while Jipujij-
kuei Kuespem is in a different ecoregion. Salmon-
ier and Fitzgerald’s Pond are on the Avalon (albeit
in different ecoregions) and so are somewhat
isolated from the other three by a narrow isthmus.
Sorensen-Dice similarity coefﬁcients do not follow
ecoregion patterns or geographic proximity (see
Table 2 and Fig. 1 for geographic relationships).
LANDSCAPE-LEVEL RELATIONSHIPS TO LICHEN DI-
. With only ﬁve protected areas, we do not
have sufﬁcient power for a statistical test of our
hypotheses that lichen diversity is related to park
size or diversity of land cover types. Although the
largest protected area, Salmonier (14.5 km
the highest reported lichen richness (144 species);
the other protected areas have richness that does
not align with their size. The smallest provincial
park, Fitzgerald’s Pond (1.9 km
) has similar
reported richness as Jipujijkuei Kuespem, which
is more than three times as large, and Fitzgerald’s
Pond has higher richness (63 species) than J.T.
Cheeseman (44 species), which is slightly larger.
Nor does there appear to be any pattern with
respect to landscover diversity, as measured by the
EOSD data. All ﬁve protected areas had very
similar landscape level diversity (between 12 and
14 EOSD land cover types) and the park with the
highest Shannon landscape diversity (Sandbanks)
did not have the highest reported lichen richness.
As noted above with respect to Sorensen-Dice
similarity in species composition, there was no
pattern where geographically close protected areas
had higher similarity. The farthest east protected
area (Salmonier) had the highest reported lichen
richness (144) and the farthest west protected area
(J.T. Cheeseman) had the lowest (44), but the other
three protected areas all had similar reported lichen
FIG. 2. The two thalli of Erioderma pedicellatum
discovered in Salmonier Nature Park. (A) Habitat.
(B,C) The thalli occur on the corresponding ﬂagged
trees directly above each image. (B) Mature thallus,
scale bar ¼1.5 cm, McMullin 16217 (observation
only). (C) Immature thallus, scale bar ¼1 cm,
McMullin 16218 (observation only).
Table 2. Summary of land cover richness (quantiﬁed as number of Earth Observation for Sustainable
Development [EOSD] land cover classes) and Shannon landscape diversity index in ﬁve parks in
Newfoundland, compared with lichen species richness.
from Salmonier (km)
No. of EOSD
Fitzgerald’s Pond 1.9 35 63 14 1.896
J.T. Cheeseman 2.4 450 44 12 1.932
Jipujijkuei Kuespem 6.8 185 61 13 1.718
Salmonier 14.5 0 144 14 1.745
Sandbanks 2.3 330 63 13 2.077
362 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 144
richness (61–64) and did not follow any west-to-
ANNOTATED SPECIES LIST.
The list is arranged alphabetically by genus and
Nomenclature follows the 20th edition of the
North American Lichen Checklist (Esslinger
2015). Any deviation from this list is the
opinion of the authors.
Species authors are cited following Brummitt
and Powell (1996), authors after 1996 follow the
20th edition of the North American Lichen
Checklist (Esslinger 2015).
Collection numbers belong to the ﬁrst author
unless otherwise stated.
Nonbolded species were not collected during the
Roman numerals correspond to collection sites
in Table 1.
Herbarium acronyms follow Index Herbariorum
*¼New species to the island of Newfoundland.
** ¼New species to the province of Newfound-
land and Labrador.
†¼Nonlichenized fungi traditionally treated
Alectoria sarmentosa (Ach.) Ach. – Corticolous
(Abies balsamea, Larix laricina (Du Roi) K.
Koch). 16003 (IV), 16091 (IX).
Arctoparmelia centrifuga (L.) Hale – Saxicolous.
16172 (VI), 16180 (VI).
Arctoparmelia incurva (Pers.) Hale – Saxicolous.
Baeomyces rufus (Huds.) Rebent. – Terricolous.
Biatora pycnidiata Printzen & Tønsberg – Cortico-
lous. R.C. Harris 53963 (I) (NY).
Bryoria americana (Motyka) Holien – Cortico-
lous (Abies balsamea, Larix laricina), terrico-
lous. 16089 (XI), 16136 (V), 16154 (VI), 16164
Bryoria bicolor (Ehrh.) Brodo & D. Hawksw. –
Terricolous (thin soil over rock). 16056 (X).
Bryoria furcellata (Fr.) Brodo & D. Hawksw. –
Corticolous (Larix laricina). 16005 (IV), 16090
(VI), 16196 (VI).
Bryoria fuscescens (Gyeln.) Brodo & D.
Hawksw. – Corticolous (Abies balsamea,Larix
laricina,Picea mariana (Mill.) Britt., Sterns &
Poggenb). 16023 (I), 16093 (XI), 16097 (V),
16131 (V), 16197 (I).
Bryoria nadvornikiana (Gyeln.) Brodo & D.
Hawksw. – Corticolous (Abies balsamea,Bet-
ula papyrifera, Larix laricina). 16000 (I), 16124
(VII), 16178 (VI).
Bryoria trichodes (Michaux) Brodo & D.
Hawksw. – Corticolous (Abies balsamea, Abies
balsamea snag). 16001 (I), 16121 (V), 16139
Calicium abietinum Pers. – Lignicolous (snag).
Calicium lenticulare Ach. – Lignicolous (snag).
16021 (I), 16084 (IX).
Caloplaca holocarpa (Hoffm. ex Ach.) M. Wad –
Saxicolous. 16129 (V).
Candelariella vitellina (Hoffm.) Mu¨ ll. Arg. –
Saxicolous. 16130 (V).
Cetraria islandica subsp. crispiformis (R¨
arnefelt – Terricolous. 16149 (VI).
Cetraria aculeata (Schreb.) Fr. – Terricolous. S.R.
Clayden 13985 (II) (NBM).
Cetraria muricata (Ach.) Eckfeldt – Terricolous.
16061 (X), 16144 (VI), 16153 (VI).
Chaenotheca balsamconensis J.L. Allen &
McMullin – Fungicolous (Trichaptum abieti-
num (Pers. ex J.F. Gmel.) Ryvarden). 16030 (I).
Chaenotheca brunneola (Ach.) Mull Arg. –
Lignicolous (snag). 16076 (IX).
Cladonia arbuscula (Wallr.) Flot. – Terricolous.
Cladonia boryi Tuck. – Terricolous. 16177 (VI),
Cladonia cenotea (Ach.) Schaer. – Terricolous.
Table 3. Sorensen-Dice coefﬁcient of lichen species similarity between provincial protected areas. Higher
coefﬁcient indicates species composition overlaps more. Total lichen species richness is given in Table 2.
Fitzgerald’s Pond J.T. Cheeseman Jipujijkuei Kuespem Salmonier Sandbanks
Fitzgerald’s Pond 1 0.5047 0.5000 0.4638 0.4921
J.T. Cheeseman 0.5047 1 0.4000 0.3511 0.4112
Jipujijkuei Kuespem 0.5000 0.4000 1 0.33317 0.4193
Salmonier 0.4638 0.3511 0.3317 1 0.4155
Sandbanks 0.4921 0.4112 0.4193 0.4155 1
2017] MCMULLIN AND WIERSMA: LICHENS OF SALMONIER NATURE PARK 363
Cladonia chlorophaea (Fl¨
orke ex Sommerf.)
Spreng. – Lignicolous (log). 16029 (I), 16162
Cladonia coccifera (L.) Willd. – Terricolous.
Cladonia cornuta subsp. groenlandica (E. Dahl)
Ahti – Terricolous. 16042 (II).
Cladonia crispata (Ach.) Flot. – Lignicolous
(stump), terricolous. 16034 (II), 16086 (VI),
16111 (XI), 16142 (I).
Cladonia cristatella Tuck. – Lignicolous (log).
Cladonia digitata (L.) Hoffm. –Lignicolous
(stump). 16032 (I).
Cladonia ﬁmbriata (L.) Fr. – Terricolous. 16044
Cladonia gracilis subsp. elongata (Jacq.) Vain. –
Terricolous. 16143 (VI).
Cladonia gracilis subsp. gracilis (L.) Willd. –
Terricolous. 16055 (X), 16184 (VI).
Cladonia grayi G. Merr. ex Sandst. – Terricolous.
R.C. Harris 53932 (I) (NY).
Cladonia macilenta var. bacillaris (Genth)
Schaer. – Terricolous. 16043 (II).
Cladonia maxima (Asahina) Ahti – Terricolous.
Cladonia multiformis G. Merr. – Terricolous.
Cladonia ochrochlora Fl¨
orke – Lignicolous (log).
Cladonia pleurota (Fl¨
orke) Schaer. – Lignicolous
(log). 16066 (IX), 16112 (XI).
Cladonia pyxidata (L.) Hoffm. – Terricolous.
Cladonia rangiferina (L.) F.H. Wigg. – Terrico-
lous. 16048 (II).
Cladonia scabriuscula (Delise) Nyl. – Terricolous.
S.R. Clayden 13991 (II) (NBM).
Cladonia squamosa Hoffm. – Lignicolous
(stump). 16033 (II).
Cladonia stellaris (Opiz) Pouzar & Vˇ
Terricolous. 16057 (II).
Cladonia stygia (Fr.) Ruoss – Terricolous. 16047
Cladonia subulata (L.) F. H. Wigg. – Lignicolous
(log). 16137 (VII).
Cladonia terrae-novae Ahti – Terricolous. 16050
Cladonia uncialis subsp. biuncialis (Hoffm.) M.
Choisy – Terricolous. 16049 (II), 16216 (VI).
Cladonia verticillata (Hoffm.) Schear. – Terrico-
lous. 16046 (II).
Cladonia wainioi Savicz – Terricolous. 16141
Coccocarpia palmicola (Spreng.) Arv. & D.J.
Galloway –Corticolous(Abies balsamea).
16072 (IX), 16119 (XI).
†Cystobasidium hypogymniicola Diederich &
Ahti – Lichenicolous (Hypogymnia incurvoides
Rass., Hypogymnia physodes (L.) Nyl.). 16104
(XI), 16161 (VIII).
Dermatocarpon luridum (With.) J.R. Laundon –
Saxicolous. 16109 (XII), 16126 (V).
Dibaeis baeomyces (L. f.) Rambold & Hertel –
Terricolous. 16045 (II), 16211 (VI).
**Ephebe hispidula (Ach.) Horw. – Saxicolous.
Erioderma pedicellatum (Hue) P.M. Jørg. –
Corticolous (Abies balsamea). 16217, 16218
Fuscidea recensa var. arcuatula (Arnold) Fry-
day – Saxicolous. 16186 (VII).
Fuscopannaria ahlneri (P.M. Jørg.) P.M. Jørg. –
Corticolous (Abies balsamea). 16117 (XI).
Graphis scripta (L.) Ach. – Corticolous (Betula
alleghaniensis,Picea mariana). 16026 (II),
Hypogymnia hultenii (Degel.) Krog – Corticolous
(Picea mariana). 16035 (IV).
Hypogymnia incurvoides Rass. – Corticolous
(conifer snag). 16053 (II).
Hypogymnia physodes (L.) Nyl. – Corticolous
(Larix laricina, Picea mariana). 16004 (IV),
Hypogymnia tubulosa (Schaer.) – Corticolous
(Picea mariana). 16010 (IV).
Hypogymnia vittata (Ach.) Parrique – Cortico-
lous (Picea mariana). 16014 (I).
Icmadophila ericetorum (L.) Zahlbr. – Lignico-
lous (stump). 16016 (I).
Imshaugia aleurites (Ach.) S.F. Mey. – Lignico-
lous (snag). 16065 (IX).
Lasallia papulosa (Ach.) Llano – Saxicolous.
16138 (V), 16171 (VI), 16181 (VI).
Lecanactis abietina (Ach.) K¨
orb. – Corticolous
(Betula alleghaniensis). 16194 (IX).
Lecanora polytropa (Ehrh.) Rabenh. – Saxico-
lous. 16142 (VI).
Lecanora muralis (Schreb.) Rabenh. – Saxico-
lous. 16122 (V).
Lecanora symmicta (Ach.) Ach. – Lignicolous
(snag). 16106 (IX).
Lecidea albofuscescens Nyl. – Corticolous. R.C.
Harris 53955 (I) (NY).
364 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 144
Lecidea tessellata Fl¨
orke – Saxicolous. 16193
Lepraria ﬁnkii (B. de Lesd.) R. C. Harris –
Corticolous (Betula alleghaniensis). 16095 (IX).
Lichinodium sirosiphoideum Nyl. – Corticolous
(Abies balsamea). 16120 (XI).
Lichenomphalia umbellifera (L. : Fr.) Redhead,
Lutzoni, Moncalvo & Vilgalys – Lignicolous
(log). 16077 (IX).
Lopadium disciforme (Flot.) Kullh. – Corticolous
(Abies balsamea, Betula alleghaniensis). 16028
(II), 16107 (IX).
Loxospora cismonica (Beltr.) Hafellner – Corti-
colous (Abies balsamea). 16025 (II).
Loxospora elatina (Ach.) A. Massal. – Cortico-
lous (Picea mariana). 16027 (II).
Melanelia hepatizon (Ach.) A. Thell – Saxico-
lous. 16173 (VI), 16179 (VI).
Melanelia panniformis (Nyl.) Essl. – Saxicolous.
Menegazzia subsimilis (H. Magn.) R. Sant. –
Corticolous. R.C. Harris 53924 (I) (NY), R.C.
Harris 53963-A (I) (NY).
Menegazzia terebrata (Hoffm.) A. Massal. –
Corticolous (Abies balsamea). 16071 (IX),
Micarea peliocarpa (Anzi) Coppins & R. Sant. –
Lignicolous (log). 16079 (IX).
**†Muellerella lichenicola (Sommerf. ex Fr.)
D.Hawksw. – Lichenicolous (Mycoblastus san-
guinarioides, M. sanguinarius). 16147 (I),
Mycoblastus afﬁnis (Schaer.) Schauer. – Cortico-
lous (Abies balsamea). S.R. Clayden 14011 (II)
Mycoblastus caesius (Coppins & P. James)
Tønsberg – Corticolous (Picea mariana).
**Mycoblastus sanguinarioides Kantvilas – Cor-
ticolous (coniferous snag). 16036 (I).
Mycoblastus sanguinarius (L.) Norman – Lig-
nicolous (snag). 16190 (VI), 16213 (X).
†Mycocalicium subtile (Pers.) Szatala – Lignic-
olous (snag). 16083 (IX).
Ochrolechia androgyna (Hoffm.) Arnold –
Corticolous (Abies balsamea). 16018 (I),
16099 (IX), 16110 (I).
Ochrolechia frigida (Sw.) Lynge – Terricolous.
16127 (VI), 16201 (VI).
Ophioparma ventosa (L.) Norman – Saxicolous.
Orphniospora moriopsis (A. Massal.) D.
Hawksw. – Saxicolous. 16185 (VI).
Parmelia saxatilis (L.) Ach. – Saxicolous. 16074
(X), 16176 (VI).
Parmelia squarrosa Hale – Corticolous (Picea
mariana). 16019 (I).
Parmelia sulcata Taylor – Saxicolous. 16060
Parmeliella parvula P. M. Jørg. – Corticolous
(Abies balsamea). 16080 (IX), 16118 (XI).
Parmeliopsis capitata R.C. Harris – Corticolous
(Picea mariana). 16012 (IV).
Parmeliopsis hyperopta (Ach.) Arnold – Cortico-
lous (Picea mariana). 16013 (IV).
Peltigera canina (L.) Willd. – Terricolous. 16150
Peltigera hymenina (Ach.) Delise – Terricolous
Peltigera membranacea (Ach.) Nyl. – Terricolous.
16103 (III), 16134 (III), 16146 (III).
Peltigera polydactylon (Neck.) Hoffm. – Cortico-
lous (Abies balsamea), terricolous. 16098 (V),
Peltigera ponojensis Gyeln. – Terricolous. 16158
Peltigera scabrosa Th. Fr. – Terricolous. 16128
†Phaeocalicium compressulum (Nyl. ex Szatala)
A.F.W. Schmidt – Corticolous (Alnus viridus
(Chaix) DC. ssp. crispa (Aiton) Turrill). 16039
†Phaeocalicium matthewsianum Selva & Tibell
– Corticolous (Betula alleghaniensis). 16040
*Phaeophyscia ciliata (Hoffm.) Moberg – Bryic-
olous. 16088 (XII).
**Placynthium ﬂabellosum (Tuck.) Zahlbr. –
Saxicolous. 16073 (X).
Platismatia glauca (L.) Culb. & C. Culb. –
Corticolous (conifer snag). 16024 (I).
Platismatia norvegica (Lynge) Culb. & C. Culb.
– Corticolous (Picea mariana). 16017 (I).
Porpidia ﬂavicunda (Ach.) Gowan – Saxicolous.
Porpidia macrocarpa (DC.) Hertel & A. J.
Schwab – Saxicolous. 16081 (IX).
Pycnothelia papillaria Dufour – Terricolous.
16115 (VI), 16133 (VI), 16200 (VI).
Ramalina dilacerata (Hoffm.) Hoffm. – Cortico-
lous (Larix laricina). 16007 (IV).
Ramalina roesleri (Hochst. ex Schaer.) Hue –
Corticolous (Larix laricina). 16006 (IV).
2017] MCMULLIN AND WIERSMA: LICHENS OF SALMONIER NATURE PARK 365
Rhizocarpon geographicum (L.) DC. – Saxico-
lous. 16195 (VI).
Rhizocarpon reductum Th. Fr. – Saxicolous.
Sphaerophorus fragilis (L.) Pers. – Saxicolous.
16062 (X), 16114 (VI), 16182 (VI).
Sphaerophorus globosus (Huds.) Vain. – Cortico-
lous (Betula alleghaniensis, Picea mariana).
16015 (I), 16212 (IX).
Stereocaulon dactylophyllum Fl¨
orke – Saxico-
lous. 16063 (III).
Stereocaulon paschale (L.) Hoffm. – Saxicolous.
*Stereocaulon subcoralloides (Nyl.) Nyl. – Sax-
icolous. 16152 (VI).
Stereocaulon vesuvianum Pers. – Saxicolous.
16148 (VI), 16157 (XII).
Thelotrema lepadinum (Ach.) Ach. – Corticolous
(Betula alleghaniensis, snag). 16054 (IX),
Trapeliopsis granulosa (Hoffm.) Lumbsch –
Terricolous. 16067 (IX).
Tremolecia atrata (Ach.) Hertel – Saxicolous.
Tuckermanopsis americana (Spreng.) Hale –
Corticolous (Betula alleghaniensis,Larix larici-
na). 16008 (IV), 16052 (II), 16116 (XI), 16155
Umbilicaria deusta (L.) Baumg. – Saxicolous.
Umbilicaria hyperborea (Ach.) Hoffm. – Saxic-
olous. 16167 (VI).
Umbilicaria muehlenbergii (Ach.) Tuck. – Sax-
icolous. 16183 (VI).
Umbilicaria polyphylla (L.) Baumg. – Saxicolous.
Umbilicaria torrefacta (Lightf.) Schrad. – Saxic-
olous. 16113 (VI), 16168 (VI).
Usnea dasopoga (Ach.) Nyl. – Corticolous (Abies
balsamea). 16082 (IX), 16199 (I).
**Usnea ﬂammea Stirt. – Corticolous (Betula
alleghaniensis). 16085 (IX).
Usnea longissima Ach. – Corticolous (Picea
mariana). 16011 (IV).
Variolaria amara Ach. – Corticolous (conifer
snag). 16022 (I). 16123 (IX).
Vulpicida pinastri (Scop.) J.-E. Mattsson & M.J.
Lai – Corticolous (Picea mariana). 16009 (IV).
**Xanthoparmelia angustiphylla (Gyeln.) Hale –
Saxicolous. 16094 (V).
Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale
– Saxicolous. 16058 (X).
Xanthoria elegans (Link) Th.Fr.’ – Saxicolous.
Xanthoria polycarpa (Hoffm.) Rieber – Saxico-
lous. 16125 (V).
Xylographa opegraphella Nyl. – Lignicolous
(snag). 16108 (IX).
Xylographa parallela (Ach.: Fr.) Fr. – Lignico-
lous (log). 16087 (V).
Discussion. Beta diversity of lichens in these
ﬁve protected areas in Newfoundland does not
follow predictable patterns. Parks in the same
ecoregion or in close geographic proximity do not
have higher similarity to each other than to parks
that are farther apart or in different ecoregions.
Across all parks, total lichen diversity (gamma
diversity) is 203 species.
The differences observed in total species
richness (alpha diversity) within parks might also
be a result of different survey efforts, and may be
affecting the lack of any predictable pattern in beta
diversity. McCarthy, Driscoll, and Clayden (2015)
spent 2 days in each park and acknowledge that
their surveys were limited to more accessible areas
and that their data should be viewed as prelimi-
nary. Similarly, we did not cover all areas of
Salmonier Nature Park (see Fig. 1) and there are at
least seven species previously collected in Sal-
monier that we did not observe. We did spend the
equivalent of 4 days surveying, twice that of
McCarthy, Driscoll, and Clayden (2015). Lichen
richness was shown to positively correspond with
survey effort by McMullin et al. (2014) in the
Arboretum in Guelph, Ontario, where they contin-
ued to ﬁnd additional species in the park after 46
collecting trips. Thus, it is reasonable to predict
higher alpha diversity in the Newfoundland
provincial parks than reported by McCarthy,
Driscoll, and Clayden (2015).
Lichen richness in Salmonier appears to be high
compared to other provincial protected areas in
Newfoundland. We found approximately twice the
number of species in Salmonier as McCarthy,
Driscoll, and Clayden (2015) did in any of their
ﬁve parks. Increased survey effort in Salmonier
may play a role, or this might be a consequence of
species-area relationships, as Salmonier is much
larger than the other parks. However, the relation-
ship between park size and lichen richness in the
other four parks does not follow any predictable
pattern, albeit with insufﬁcient statistical power (n
¼5) for any robust test. Our analysis of habitat
366 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 144
diversity does not suggest that Salmonier has
dramatically higher habitat diversity, at least not as
measured using the EOSD data. However, EOSD
is based on satellite-based remote sensing, and thus
does not delineate ﬁne-scale habitat types, which
are important for lichens. The much larger size of
Salmonier means that it is possible that there is a
higher diversity of meso- and microhabitats
compared to the other parks surveyed by Mc-
Carthy, Driscoll, and Clayden (2015). We can
speculate that Salmonier might have uniquely high
lichen diversity compared to the other four
provincial parks. This could be due to simple
species-area relationships, a higher diversity of
meso- and microhabitats, the fact that this part of
the province has older forest stands than most of
the other regions, unique climate patterns, or some
combination of these factors. However, we do not
have data suitable to test these hypotheses.
One species discovered at Salmonier, Erioderma
pedicellatum (Fig. 2), is a foliose cyanolichen with
convex apothecia that are red-brown to brown,
conspicuous hairs on the upper surface, and no
vegetative propagules (Galloway and Jørgensen
1987, Maass and Yetman 2002). It is listed as a
species at risk of extinction at several levels.
Provincially, it is listed as ‘‘vulnerable’’ by the
Newfoundland and Labrador Department of Envi-
ronment and Conservation (2016). Federally, the
Committee on the Status of Endangered Wildlife in
Canada has listed it as a species of ‘‘special
concern’’ (COSEWIC 2014). Globally, it is listed
by the International Union for Conservation of
Nature as ‘‘critically endangered’’ (Scheidegger
2003). Two thalli were discovered in the moraine
and bog ecosystem in the southwest corner of the
park. We expect that further survey efforts in this
area will result in additional thalli being discovered.
In 2000, thalli of E. pedicellatum were transplanted
onto four balsam ﬁr trees in the park along the
wildlife boardwalk near the current great horned
owl (Bubo virginianus) enclosure by Christoph
Scheidegger and Mac Pitcher (M. Pitcher, retired.
Former employee of Salmonier Nature Park). We
extensively searched the area, but the transplants
appear to have been unsuccessful.
Several lichens and one lichenicolous fungus
discovered in Salmonier are also uncommon or
new to Newfoundland and Labrador. Six species
are reported for the ﬁrst time in the province:
Ephebe hispidula, Muellerella lichenicola, Myco-
blastus sanguinarioides, Placynthium ﬂabellosum,
Usnea ﬂammea, and Xanthoparmelia angustiphyl-
la. Ephebe hispidula is a ﬁlamentous cyanolichen
that was found growing on a boulder in moving
water. It is distinguished from the more common
Ephebe lanata (L.) Vain. by the presence of spine-
like perpendicular branchlets along the main
branches (Henssen 1963). Muellerella lichenicola
is a lichenicolous fungus that was regularly
observed growing on the thalli of Mycoblastus
sanguinarioides and M. sanguinarius. It is distin-
guished by its small (,125 lm in diameter) dark
and immersed to sessile perithecia with pale brown
two-celled spores (Triebel and Kainz 2004).
Mycoblastus sanguinarioides was recently report-
ed from northeastern North America by Spribille,
Klug, and Mayrhofer (2011). They show that
specimens in this region previously identiﬁed as M.
sanguinarius might be M. sanguinarioides,a
similar species that was only known from the
Southern Hemisphere. Mycoblastus sanguinar-
ioides differs in having birefringent crystals in the
hymenium that do not occur in M. sanguinarius
(Spribille, Klug, and Mayrhofer 2011). Pla-
cynthium ﬂabellosum is a subfoliose cyanolichen
on boulders in moving water. It differs from other
species of Placynthium by its overlapping, radially
extended lobes with crenulate margins, its lack of a
hypothallus, and its occurrence on noncalcareous
rock (Henssen 1963, Schultz 2002). Usnea ﬂam-
mea is a tufted fruticose lichen that was found
growing on Betula alleghaniensis. Among the
species of Usnea in Newfoundland and Labrador it
can be identiﬁed by its pale base, pale to white
medulla and cental axis, and the presence of
soredia, isidia, and stictic and menegazziaic acids
(KOHþyellow to orange-red) (Hinds and Hinds
2007). Xanthoparmelia angustiphylla was grow-
ing on lakeside boulders. It is distinguished from
other species of Xanthoparmelia by a lack of
vegetative propagules, a black lower surface, and
the presence of stictic and nortstic acids (KOHþ
yellow to orange-red) (Hale 1990, Thomson 1993).
Two additional species are reported for the ﬁrst
time from the island of Newfoundland: Phaeophy-
scia ciliata and Stereocaulon subcoralloides.
Phaeophyscia ciliata was growing on bryophytes
on lakeside boulders. It is distinguished from
similar species in the province by a lack of
atranorin in the upper cortex (KOH), a black
lower surface, the absence of vegetative propa-
gules, and lobes that are ,1 mm wide (Moberg
1977, Esslinger 1978). It was previously reported
2017] MCMULLIN AND WIERSMA: LICHENS OF SALMONIER NATURE PARK 367
in the province from southeastern Labrador (Lynge
1947). Stereocaulon subcoralloides is an erect
fruticose species that was also growing on lakeside
boulders. It differs from other species of Stereo-
caulon by its cylindrical coralloid phyllocladia that
are ,0.6 mm long, pseudopodetia without
tomentum, and the absence of stictic and norstic
acids (Lamb 1978, Hinds and Hinds 2007). It was
previously reported in the province from western
Labrador by Lamb (1977) and Thomson (1984).
Our results provide a better understanding of the
distribution of lichens in Newfoundland and
Labrador and within the Avalon Forest Ecoregion.
These baseline data aid our ability to analyze
landscape diversity patterns for lichens in the
province and in the development of the required
evidence for sound conservation strategies (Suther-
land et al. 2004). Within Salmonier, rare species,
new species to the park, and rare habitats can now be
better acknowledged. Changes to the lichen biota in
the park can now be monitored as well. Compre-
hensive surveys of spatially deﬁned areas or
ecosystems in Newfoundland and Labrador, how-
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