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A geographical analysis of alpine lichen in Rocky Mountain National Park

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Title:
A geographical analysis of alpine lichen in Rocky Mountain National Park
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Ahl, Erik Douglas ( author )
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Denver, Colo.
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University of Colorado Denver
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Master's ( Master of science)
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University of Colorado Denver
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Department of Geography and Environmental Sciences, CU Denver
Degree Disciplines:
Environmental sciences

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Lichens ( lcsh )
Lichens ( fast )
Rocky Mountain National Park (Colo.) ( lcsh )
Colorado -- Rocky Mountain National Park ( fast )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

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Review:
In high alpine ecosystems, landscapes often seem barren to the naked eye due to intense exposure from UV light, temperature fluxes, and desiccation. Species present are only those resilient enough to have adapted to such harsh conditions, and these are usually ground-hugging species such as those in Biological Soil Crusts (BSCs). Consisting of various species of cyanobacteria, algae, lichens, and mosses, BSCs have been studied for decades in dry land environments all over the world, yet little BSC research focuses on high altitude ecosystems. This study set out to analyze BSC distribution and characteristics. In situ, however, lichens dominated rocks and soil at the chosen sites and thus became the focus for the study. The four sites (three high alpine, one subalpine) utilized were already under study in Rocky Mountain National Park (RMNP) with the Rocky Mountain Inventory and Monitoring Network (ROMN) in conjunction with the Global Observation Initiative in Alpine Environments (GLORIA). Implementing field techniques outlined by GLORIA, species were photographed, identified, inventoried and then analyzed using SPSS statistical software. It was suspected that findings would show correlations between aspects, soil, and rock types. Relationships between some species and specific rock types, such as siliceous and granitic, were initially discovered, as well as variability in types among other species.
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Includes bibliographical references.
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System requirements: Adobe Reader.
Statement of Responsibility:
by Erik Douglas Ahl.

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Auraria Library
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987265725 ( OCLC )
ocn987265725
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LD1193.L547 2016m A55 ( lcc )

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Full Text
A GEOGRAPHICAL ANALYSIS OF ALPINE LICHEN IN ROCKY MOUNTAIN
NATIONAL PARK by
ERIK DOUGLAS AHL B.S., Western Carolina University, 2011
A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirements for the degree of Master of Science Environmental Sciences program
2016


2016
ERIK DOUGLAS AHL ALL RIGHTS RESERVED
11


This thesis for the Master of Science degree by Erik Douglas Ahl has been approved for the Environmental Sciences Program by
Casey Allen, Chair Frederick Chambers David Knochel
December 17th, 2016
in


Ahl, Erik Douglas (MS, Environmental Sciences)
A Geographical Analysis of Alpine Lichen in Rocky Mountain National Park Thesis directed by Professor Casey Allen
ABSTRACT
In high alpine ecosystems, landscapes often seem barren to the naked eye due to intense exposure from UV light, temperature fluxes, and desiccation. Species present are only those resilient enough to have adapted to such harsh conditions, and these are usually ground-hugging species such as those in Biological Soil Crusts (BSCs). Consisting of various species of cyanobacteria, algae, lichens, and mosses, BSCs have been studied for decades in dry land environments all over the world, yet little BSC research focuses on high altitude ecosystems. This study set out to analyze BSC distribution and characteristics. In situ, however, lichens dominated rocks and soil at the chosen sites and thus became the focus for the study. The four sites (three high alpine, one subalpine) utilized were already under study in Rocky Mountain National Park (RMNP) with the Rocky Mountain Inventory and Monitoring Network (ROMN) in conjunction with the Global Observation Initiative in Alpine Environments (GLORIA). Implementing field techniques outlined by GLORIA, species were photographed, identified, inventoried and then analyzed using SPSS statistical software. It was suspected that findings would show correlations between aspects, soil, and rock types. Relationships between some species and specific rock types, such as siliceous and granitic, were initially discovered, as well as variability in types among other species.
iv


The form and content of this abstract are approved. I recommend its publication
Approved: Casey Allen


TABLE OF CONTENTS
CHAPTER 1......................................................................1
I. INTRODUCTION..............................................................1
1.1 Internal Function......................................................2
1.2 Classification.........................................................2
1.3 Effects on Environment.................................................4
1.4 Lichen Distribution....................................................5
CHAPTER II.....................................................................8
II. .SITE SETTING............................................................8
CHAPTER III...................................................................10
III. METHODS................................................................10
3.1 Field Work............................................................10
3.2 Post Field Work.......................................................11
3.3 Statistical Analysis..................................................13
CHAPTER IV....................................................................14
IV. RESULTS................................................................14
4.1 VQS...................................................................14
4.2 PIK...................................................................15
4.3 GLA...................................................................16
4.4 JSM...................................................................18
4.5 Simpsons Index of Diversity..........................................20
4.6 Jaccards coefficient.................................................21
4.7 Rank Abundance........................................................23
4.8 SPSS..................................................................26
CHAPTER V.....................................................................32
V. DISCUSSION..............................................................32
CHAPTER VI....................................................................34
VI. CONCLUSION.............................................................34
REFERENCES....................................................................37
vi


APPENDIX
43
A. LICHEN LIST FROM RMNP..........................................43
B. SPECIES TABLES.................................................47
C. SITE TABLES....................................................66
D. SITE TOTALS....................................................71
E. SPECIES PHOTOGRAPHS............................................73
F. GOOGLE EARTH HISTORICAL IMAGERY................................93
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CHAPTER I
INTRODUCTION
Somewhere amongst macro and microorganisms, splayed across soil, rocks, and trees are often brightly colored plant-like creatures called lichens. From small patches to large carpet-like spreads, these often overlooked non-vascular organisms can be found throughout the world and across all ecosystems: from the hot deserts, to cold alpine tundra, rainforests, coastal systems and even Antarctica. Lichens are part of a larger collection of species, along with algae and mosses that can be classified under the umbrella of Biological Soil Crusts (BSCs) (Belnap and Lange 2003b, Belnap et al. 2001, Purvis 2000).
Though much has been discovered in recent decades regarding lichens, most studies have focused on lower elevation species, centering specifically in hyper arid and arid deserts (Allen 2005, Allen 2010). Excluding a handful of studies conducted outside of the United States (Breen and Levesque 2008, Pohl et al. 2009, Karsten, Lutz and Holzinger 2010, Yoshitake et al. 2010) little research has been conducted in regards to higher elevation lichen in the United States.
With this in mind, this study aims to shed more light on under studied alpine lichen in the United States. Taking place in Rocky Mountain National Park (RMNP), Colorado, four sites were selected as proxies for determining richness, abundance, and cover of lichen species at differing elevations, and teasing out their locational relationships in regards to aspect, elevation, and site location to investigate perceived differences between alpine and subalpine (Belnap and Lange 2003a, Brodo, Sharnoff and Sharnoff 2001b).
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First off, a brief discussion of the lichens internal function (1.1) and classification (1.2) will be offered, followed by their role in the environment (1.3). Then a description of the studys site setting (II) is offered, followed by an explanation of methods implemented in the field and for the post-field work (III). Finally, results (IV) are presented and analyses are outlined along with an in-depth discussion (V) of those techniques and results leading to a brief conclusion (VI).
1.1 Internal Function
Though numerous vascular and nonvascular plant species inhabit these high elevation landscapes, the predominant species found sprawling across the rocks and soils in often-bright contrast are lichens. Carl Linnaeus, the father of taxonomy, is said to have called lichens the poor trash of vegetation and assigned only one genus (Lichen) to encompass all lichens (Walewski 2007). Little did he realize that lichens are not so easily classified into any normal plant group. They represent a complex relationship developed between two separate organisms working together for a mutually beneficial relationship. This symbiosis is the marriage between a fungi and a photosynthetic partner of algae or cyanobacteria (Brodo, Sharnoff and Sharnoff 2001c), a partnership that allows for the algae to live within the body of the fungus, while the fungus consumes algal cells for its own nourishment. These algal cells replenish at a rate much quicker than they are consumed, allowing for a very efficient form of parasitism in which both parties benefit (McCune and Goward 1995).
1.2 Classification
There are approximately 14,000 lichen species throughout the world, each of which varies in shape, size, and color. Generally, lichen are classified into four growth forms
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utilizing its vegetative body known as the thallus: foliose, fruticose, crustose, and squamulose (Brodo et al. 2001c).
Foliose lichens, which encompassed 10 of the 19 species in this study, often portray a leafy in shape, as the word implies. These lobed appendages host a more or less flattened thallus and are securely anchored to the substrate through the use of hyphae. Hyphae are filamentous elements of the fungal component of the lichen, consisting of cells arranged in threads of varying shapes, sizes, and branching patterns. Also instrumental for anchorage to substrate by foliose lichens are rhizines branched or unbranched bottom-growing appendages that often occur in tufted or fibrous bundles (Brodo et al. 2001c).
Fruiticose lichens, two of the 19 species in this study, come in a more shrubby, three dimensional form with a radial symmetry in their stalks (McCune and Goward 1995). Growing in a more erect or pendent fashion, they resemble foliose lichens, but their thali are comprised in a more three- than two-dimensional fashion (Brodo et al. 2001c).
Crustose lichens resemble, as the name implies, a crust across the chosen surface substrate and were represented in this study by seven of the 19 species. Found in a vast array of radiant colors, these lichens grow on and among the particles of the substrate forming small to very large patches that can be quite thick or rough (Brodo et al. 2001c). What differentiates crustose from the foliose and fruticose is the absence of a lower cortex, or lower fungal layer.
Squamulose lichen, which were not represented in the study sites, consist of a cross between foliose and crustose in both growth and formation (Brodo et al. 2001c). These
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growth forms are composed of clusters in small roundish flakes and come in a various shapes with slightly lobate margins (McCune and Goward 1995).
1.3 Effects on Environment
Lichens play significant roles with their surrounding biota (Kar et al. 2014). For example, like most vegetative species, lichens no matter their location, from desert plains to alpine ecosystems help with C sequestration and N fixation (Belnap and Lange 2003b, Johnston 1997, Barger et al. 2006, Yoshitake et al. 2010). These types of desiccating resilient organisms are known as poikilohydric (Viles 2008), because they are able to dry out and suspend respiration without any negative side effects, while being environmentally dependent on their water (Green and Lange 1995). Exercising such environmental resilience to adverse conditions, such as those experienced by all vegetation in the high elevations of the alpine tundra in Colorado, lichen are known to live and thrive for hundreds of years (Lalley and Viles 2007). They have the ability to colonize just about anywhere including bare rocks and soil. Lending to not only fixation and influencing hydrological patterns, their influence on the erosion of soil and rock can be paramount (Zelikova et al. 2012). The hyphae appendages extending below the thallus, and being able to bore into solid rock substrate, can allow for the speeding up of local rock decay processes (Mcllroy de la Rosa, Warke and Smith 2012). This mechanical decay can also be exacerbated by chemical decay from the carbonic acid formed in lichen metabolic activities (Raggio, Green and Sancho 2015), eating away at rocks rich in calcium carbonate that they reside on (Brodo et al.
2001c).
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At lower elevations, lichen can also be found colonizing the exterior of trees along their bark (Ames et al. 2012). Faster growing foliose and fruticose species tend to colonize first, with crustose species being the last to show up on tree bark. As forest inhabitants, they affect the dispersion of rainwater nutrients as they flow down the body of the tree (Brodo et al. 2001c). This same process occurs with nutrients absorbed from fog, thus affecting what concentrations are reflected in the ground below, affecting the surrounding plant growth.
Lichen also play an important role in environmental monitoring (Zedda et al. 2011, Shukla 2014, Bosch-Roig et al. 2013) as they are well known for their sensitivity to air pollution (Corbridge and Weber 1998, Kularatne and de Freitas 2013). A sensitivity, that is directly related to their uncanny ability to absorb chemicals rapidly from the air (Brodo et al. 2001c, Belnap et al. 2001, Walewski 2007, Kularatne and de Freitas 2013). Sulfur dioxide and components of acid rain (Purvis 2000, Cuculovic et al. 2014) (sulfuric and nitric acids) are especially detrimental to lichen species. In areas of high industrial pollution, lichen tend to be absent and this can be used as a biomarker to an areas ambient pollution (Brodo, Sharnoff and Sharnoff 2001a).
1.4 Lichen Distribution
Almost as varied as their biology and structure, lichens also remain a widespread organism around the world (Feuerer and Hawksworth 2007). (Brodo et al. 2001c) notes in his book that all lichen species have climatic tolerances and physiological constraints (Peksa and Skaloud 2011, Gauslaa 2014). So, with that in mind, the authors classified distribution patterns in relation to major climatic regions across the North American continent. Their classification is broken down into eight climatic elements: Arctic Alpine, Boreal, Temperate,
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Central Grasslands, Western Sector, Western Montane, Madrean, Tropical, Oceanic and Maritime.
The Arctic-Alpine in the more northern parts of the continent is an arid ecosystem with harsh elements (Bonan 2010, King et al. 2012). Typically situated at elevations above 3,505 M, only the resilient species like lichen may thrive under the harsher elements (Bjerke 2008) experienced there including freezing temperatures, snow, high winds greater UV exposure and desiccation (Belnap 2006). Most lichen species residing in these elements are able to shut down internal processes during long dry spells, enduring the severe frost in winter, only to spring into action once the first rain or mist lays moisture on them (Dahl 1954, Stephenson 2010, Belnap 2006).
Alpine environments, as in (Fig.. 1.4.1), comprise about 20% of land cover in some areas. Within these elevations, alpine ecosystems are deemed an important source of information due to their particular vulnerability to climate change and lack of environmental studies on their ecophysiological performance (Karsten and Holzinger 2012, Belnap and Lange 2003a, Breen and Levesque 2008). With this in mind, this lichen study was focused on these important ecosystems found within Rocky Mountain National Park, and how species abundance and diversity might contrast to a subalpine site.
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Alpine Tundra
Figure 1.4.1 Alpine Tundra locations around the world (dark blue), typically above 3,300 meters, credit https://infogr.am/alpine-9177284109
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CHAPTER II
SITE SETTING
Under the scope of few studies in relation to alpine environments, as well as, the lack of lichen based studies in alpine systems, the author has engaged in a study pertaining to alpine lichens in RMNP. This studys intention rests in teasing out locational relationships of lichens while building a biological inventory to act as baseline data for further studies, while adding to previous in situ inventories. All work is in accordance with four previously studied sites under the parks Rocky Mountain Monitoring Network (ROMN) and Global Observation Research Initiative in Alpine Environments (GLORIA).
Figure 2.1 Satellite image of R.M.N.P. displaying the location of the four study sites.
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ROMN is comprised of 32 vital monitoring sites spread throughout the National Park Service under program manager Mike Britton. In conjunction with GLORIA, an international monitoring network established in 2001, ROMN established these sites on RMNP in 2009 (Ashton 2011). GLORIA summit criteria, adopted for use by ROMN, outlined in their field manual (Pauli et al. 2004) was the basis for site choice. Located in the higher elevations of RMNP, GLORIA had three unnamed alpine sites and one subalpine already under monitoring (Table 2.1): (VQS, PIK, GLA) and Jack Straw Mountain (JSM), respectively. These sites are located in the southwestern part of the park by Milner Pass and accessed via the Ute to the Mt. Ida trail. The three alpine sites run along the continental divide in a north (VQS) to south direction, with GLA acting as a southern terminus. The subalpine site (JSM) sits just west of the ridge to Mt. Ida and was accessed by parking at the Lake Irene trailhead, crossing trail ridge road, and bush whacking to the summit. The alpine and subalpine are typically ecosystems that are found above tree line, though this elevation changes with increased latitude. In Colorado, the elevation range is approximately 3,048 m to 3505.2 m for subalpine and 3505.2 m and higher for an alpine designation. GLORIA also calls for the sites to be similar in disturbance, in this case meaning limited park traffic and geology. All four sites have similar geology consisting of granite, gneiss, and schist, with a couple instances of quartz outcrops (United States Department of Agriculture 2007).
Table 2.1 Name, location, elevation, and vegetation zones of the four sites (Ashton, 2011).
Gloria Summit Code Summit Name Latitude (degrees) Longitude (degrees) Elevation (M) Vegetation Zone
GLA Shoulder of Mount Ida 40.380617 405.770072 3862 Upper alpine
PIK Unnamed 40.392956 405.790538 3715 Lower/upper alpine
VQS Unnamed 40.406275 405.799238 3623 Lower alpine
JSM Jackstraw 40.394304 405.813083 3520 Lower Alpine/subalpine
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CHAPTER III
3.1 Field Work
METHODS
So as not to reinvent the wheel and stay in accordance to practices implemented by GLORIA and ROMN, monitoring protocols developed by GLORIA were utilized for this study. As Ashton (2001) asserts:
The goals of the GLORIA program are to provide a global baseline for vegetation monitoring in alpine environments to assess the risks of biodiversity loss and ecosystem instability from climate change. The methodology is extended by cooperators, such as the ROMN, to create a long-term monitoring network at the global scale.
Although unnecessary for this studys purpose, Ashton (2011), also notes that ROMN contains components for soil condition, tree line movement, and human disturbance.
The GLORIA design calls for four sentinel sites, sites that are all similar in geology, climate, disturbance, and land-use history, such as the four used in this study. Eight summit area sections are used to measure the dimensions of the peak (area and slope) from the highest point to 10m below (Ashton 2011). GLORIA uses this method to measure ground cover that has been divided into seven classes. However, it is in the interest of this study to do a biological inventory, so quadrat clusters will be placed within the plots in 3m sections, equating to 8 quadrats per peak placed in respect to aspect. This is a GLORIA standard to measure vegetation change (cf., Ashton, 2011) and will be implemented by this study for inventorying and noting disturbance effects to recorded species of lichen.
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Summit coordinates were supplied by GLORIA and entered into a Garmin GPS 60CSx handheld GPS unit to ensure correct site location. Each summit has been marked with a pile of rocks that also served as protection for their data collection units. 3m quadrats were measured using a Komelon 6611 100ft open reel fiberglass tape measure and staked with 6.35cm x 8.89cm x 53.34cm orange wire survey flags. These flags were temporarily placed during the duration of the study and removed upon completion of that days coverage. Flags were placed into the measured corners of each quadrat, resembling a tic-tac-toe formation. A GPS coordinate was logged with the Garmin unit at all four outer comers of each quadrat. Each of the four aspects per site was measured out into upper and lower sections. The upper section was from peak to five meters below and the lower section was between the five meter mark and 10 meters from peak. A 3m quadrat was measured out in random placement within the confines of the upper and lower measurements of each aspect. Photographs were taken using a Nikon D80 10.2 MP camera for documentation of in situ species identification. All photographic data along with site appropriate notes were recorded in a Rite in the Rain field journal and site information was written and photographed on a dry erase board.
3.2 Post Field Work
The PI and advisor reviewed the photographs and using a dichotomous key in the book Lichens of North America (Brodo et al. 2001c) identified each of the species represented. Once identified, each image was then checked against a list of species that RMNP has published on a website (Appendix A),as well as double checked in a series of other books: Macrolichens of the Northern Rocky Mountains (McCune and Goward 1995), A Color Guidebook to Common Rocky Mountain Lichens (St. Clair 1999), and A Rocky Mountain Lichen Primer (Corbridge and Weber 1998).
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In total, 19 species were positively identified (Appendix B) during the photographic review. For this study, the term species denotes an individual genus and species (eg., Acarospora contigua) and total represents the sum of a species individually, or the sum of all encompassed species in relation to a site, including aspect, slope and quadrats. Then, using data sheets and ticking off these counts, all species were tallied up per lm segment for a total of nine segments per quadrat. This was carried out for every upper and lower segment of each aspect as each site. Counts are approximations of species abundance, and the precision varied depending upon overlap of species on substrates. While multiple photographs were taken to improve visibility of counts, some photos in review had blurry edges or lighting that failed to lend a definitive identification of certain species. Only those species that could be identified through clear, focused, well-lit photographic evidence were accounted for in this study. All numbers were then tallied per species, elevation, aspect, and site, before being entered into excel spreadsheets (see Table 3.2.1 for total counts).
Table 3.2.1 Totals of species accounted for by site, upper/lower, and aspect.
VQS PIK GLA JSM
Upper Total 4804 8277 8847 6420
Lower Total 3340 4883 8456 5380
Site Totals 8144 13160 17303 11800
Overall Study Total 50407
Aspect Totals
West 2472 2710 6160 3199
South 2864 3515 6861 3428
North 1590 2889 4282 3965
East 1218 4046 N/A 1208
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3.3 Statistical Analysis
The data were analyzed in IBMs SPSS (IBMCorp. 2016) software for statistical trends, as well as assessed with non-parametric tests using the Simpsons Diversity Index (Simpson 1949), Jaccards coefficient (Stohlgren et al. 1997) in order to conduct tests of species overlap, and a ranking of species abundance. A Generalized Linear Model (GLIM) (following Bolker et al. 2008) assuming a negative binomial distribution and log link function was used to compare the count data describing lichen species abundance across sites and aspects. Site (four levels) and aspect (four levels), were treated as fixed effects. Following a statistically conservative approach, and due to a lack of a-priori hypotheses about any expected site by aspect interactions, this factor was not included in the final GLIM. The generalized linear model with a negative binomial distribution was the approach recommended for the analysis of over dispersed count data (O'Hara and Kotze 2010, Bolker et al. 2008). Post-hoc pairwise comparisons using the Bonferroni adjustment were made by site and aspect, and species counts were considered significantly different at p <0.05. Descriptive statistics of species abundance and averages across all sites, aspects, and distances to the summit (upper and lower) were also reported.
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CHAPTER IV
RESULTS
A few initial general findings stood out after the species were tallied for each of the sites (see Appendices II-IV). Across all sites, 570 individual counts of lichen abundance were conducted, with four site levels, four aspects (except for GLA), and two distances to summit (upper and lower), for a total of 30 quadrats, with 19 species observed within each. First, sites totals for the three alpine sites (VQS, PIK and GLA) will be examined. Then, the subalpine (JSM) sites totals and species counts, a discussion of the Simpsons Diversity Index and how it relates to each site follows. Next, Jaccards coefficient of species to assess species overlap across all four sites will be outlined, including a brief display of rank abundance. Finally, a review of overall statistical findings (based on data obtained using SPSS) that were derived at the site level, aspect level, and aspects within site is offered.
4.1 VOS
This site had the lowest overall total number of species at 8,144. The upper part of VQS had a total of 4,804 species, with the largest amount found on the southerly aspect (1,487). The most prevalent of species was Umbilicaria virginis at 1,380. Umbilicaria americana and Vulpicidapinastri both had zero presence. The lower portion of VQS had slightly lower numbers and had an approximate total of 3,340 species with the Southerly aspect once again exhibiting the largest numbers at (1,377). As with the upper aspect, U. virginis (752) was the most prevalent species tallied, and V. pinastri and Thamnolia subuliformis having zero presence at this distance from the summit.
14


This site was fairly homogenous in cover between metamorphic rocks and grass in both the upper and lower portions examined and across all four aspects. This is reflected in the numbers in both aspect and relative distance to summit, with the upper portion having larger numbers due to there being slightly more rocks than grass (Fig. 4.1.1).
Figure 4.1.1 Looking West to the summit from the lower East quadrat on VQS.
4.2 PIK
This site, located north along the ridge from VQS had the second largest abundance of species at 13,160 and included all 19 species that have been identified in this study. This is in large part to the greater amount of rocks found at this site in comparison to the previous VQS site (Fig. 4.2.1). The upper portion had an approximate total of 8,277 species with the
15


highest proportion being on the Easterly (2,857). Diploschistes muscorum and T. subuliformis, soil based species, had zero presence in the upper portion of this site and once again was dominated by U. virginis (1,661). The lower portion had an approximate total of 4,883, with the largest total on the North (1,489). All 19 species were accounted for on the lower portion.
Figure 4.2.1 view to the summit from the lower East quadrat at PIK.
4.3 GLA
This site was further north along the ridge and slightly East of the others, as it was situated along the shoulder of Mt. Ida, (Fig. 4.3.1). This site had the largest approximate abundance of species of all four sites with 17,303. Only three aspects were accounted for on
16


this site due to the East aspect being on the edge of a steep cliff and thus inaccessible by the PI. The upper portion assessed had a total of 8,847 species accounted for with half of that being found on the Southerly (4,131). Both species of Vulpicida had zero presence, with U. virginis dominating the rocks once again. The lower section of GLA had an approximate total of 8,456. All three aspects in the lower portion had totals that were closer to one another than the upper with the Westerly aspect hosting the most (3,343). As in the upper portion and due to the abundance of rock cover, soil based species V. pinastri, V. tilesii and T. subuliformis had zero presence. U. virginis once again dominated the rocks (3,575) closely followed by Rhizocarpon geographicum (3,504). The heavy presence of large, slabbed rock in the site (Fig. 4.3.1) seemed to affect the high numbers of rock-based species and lower numbers of soil-based species.
Figure 4.3.1 view to the North on GLA from the lower South quadrat.
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4.4 JSM
In contrast to the other three sites, JSM was situated west of the alpine ridge at a lower elevation, (Fig. 4.4.1). This site had the third highest count in relation to the other three with an approximate total of 11,800. This site was very similar to VQS in that it was a balanced mix of rock and grass, with the exception of the East aspect. The East side appeared to have had some disturbance from some point in time, which left it with little rock and grass. This was a stark contrast to the other three aspects and was reflected in the numbers. Upper JSM had a total of 6,420 species with the North aspect (Fig. 4.4.2) accounting for 1,896. East upper was the lowest count of all four with 1,131. This trend was reflected in the lower portion of the East aspect where disturbance seemed high and only accounted for a total of 77 species, the lowest of any aspect from any site in this study. This seems to be in part of the difference in ground cover. Very few rocks, little grass, and various patches of soil dominated this portion of JSM. The lower part had a species total of 5,380 with the North aspect once again exhibiting the highest number at 2,069.
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Figure 4.4.1 Google Earth satellite imagery of the four study sites in RMNP.
Figure 4.4.2 the view South from the lower North aspect on JSM.
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4.5 Simpsons Index of Diversity
For this study, the Simpsons index of diversity D = 1 (H N(^N_1^) was
implemented to test the lichen diversity within the datasets. Where n = total number of individuals of a particular species at a site (VQS, PIK, GLA, JSM) and N = total count of all species within the study sites, to characterize the richness, abundance, and evenness of lichen across sites (Arcadia 2013, Morris et al. 2014). The value of the index ranges from 0 to 1, with 0 having the lowest site diversity, and 1 the highest site diversity. The calculated Simpsons index of diversity was high across all of the sites (range was 0.860 0.884), which was to be expected with 19 species found in the alpine and subalpine, which are common species for this region and these elevations. The lowest index was at VQS (0.86) and the highest at PIK (0.884) with GLA (0.864) and JSM (0.871) falling very closely in between. The Simpsons index of diversity provided an overall estimated probability of 87% that if one were to randomly choose two lichen individuals from the overall community assessed as these sites, two different species would be picked in the sample, indicating a fairly high level of lichen diversity across sites.
Table 4.5.1 Simpsons Index per site
Site D=
VQS 0.860
PIK 0.884
GLA 0.864
JSM 0.871
20


0.89
0.885
0.88
0.875
0.87
0.865
0.86
0.855
0.85
0.845
Sites
VQS
PIK
GLA
JSM
Figure 4.5.1 A graphical representation of the Simpsons Index scores from Table 4.5.1 per site location (VQS, PIK, GLA, JSM)
4.6 Jaccards Coefficient
Jaccards coefficient, a non-parametric test to compare species overlap among sites, it was used for pairwise comparisons of observed lichen species identity overlap among sites and aspects, where / = A/{A + B + C) A = the number of species (richness) found in both of the compared sites, B = the number of species found in the first site, but not present in the second site, and C = the number of species found in the second site, but not present in the first site (Stohlgren et al. 1997, Chong and Stohlgren 2007). If Jaccards coefficient is equal to 0, then there is no overlap in species lists between sites, whereas if the coefficient is equal to 1.0, then there is complete overlap of all species across the two sites compared. Using this approach, this study first analyzed site diversity overlap using pairwise comparisons at the site level (see Table 4.6.2). Sites ranged from a low of 16 to a maximum of 19 species present (Table 4.6.1), and upon comparing sites, the Jaccards value ranged from 0.842 to
21


0.947, indicating generally high species overlap across sites (with the 2-3 soil-based lichen species commonly the ones absent, or in low abundance, across the sites except at PIK).
Next, Jaccards overlap was compared across plots that were within individual study sites (the subalpine site JSM and three alpine sites: VQS, PIK, GLA), and little variability was observed with respect to the Jaccard's coefficient in comparing overlap of within-site richness between aspects and proximity to the summit (upper and lower quadrats). Lastly, across sites, some notable differences arose when comparing aspects. The northerly and westerly aspects exhibited large overlap in Jaccard's; however, when comparing the easterly and southerly aspects of the mean of the three alpine sites together (by pooling the upper and lower species presence/absence in each site) to the subalpine sites value (also pooling the upper and lower quadrats), the degree of Jaccards overlap was substantially lower (Table 4.6.3). The observed lichen species richness was highest in the alpines easterly and southerly aspects (mean of all three alpine sites), compared with these respective aspects in the subalpine, with Jaccards at 0.62 comparing east, and 0.74 comparing south. JSM, the subalpine site, had less exposed rock cover and vegetation broken up by patches of soil or loose rock.
Table 4.6.1 Site Species values for Jaccards coefficient showing how many of the 19 species are represented per site.
VOS PIK GLA JSM
18 19 17 16
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Table 4.6.2 Jaccard's pairwise comparisons.
Site Comparisons J=
PIK 18/(18+0+1) 0.947
vos JSM 16/(16+2+1) 0.842
GLA 18/(18+1+1) 0.9

PIK GLA 17/(17+2+0) 0.894
JSM 16/(16+3+0) 0.842

GLA JSM 16/(16+1+0) 0.941
Mean ./= 0.896
Table 4.6.3 Jaccard's East and South Alpine and Subalpine comparisons.
Jaccard's Alpine East = 0.615384615
compared to subalpine east

Jaccard's Alpine South = 0.736842105
compared to subalpine south
4.7 Rank Abundance
Rank abundance curves represent a way of plotting relative species abundance to visualize the biodiversity of species richness and evenness in a study. The curve displays the proportional abundance of each species, and compares it to that species abundance rank. The curves are a simple way for one to visualize whether species assessed are common or rare across sites. Data were obtained, per site, by taking the total for each species and dividing it by the site total. Based on the resulting quotient, each was assigned a rank (Table 4.7.1) and then graphed to show the resulting curves per site (Fig. 4.7.1). For all three Alpine sites (VQS, PIK, GLA) the top ranked species was Umbilicaria virginis, while Acarospora contigua was top ranked at the subalpine site. Typically, there tend to be a few species within communities that exhibit dominance (maintain high abundance with respect to others), with
23


the remainder of species tending to be less common or rare, resulting in a dramatic drop and steep slope in species rank-abundance curves. However, in this case with alpine and subalpine lichen, there exist a substantial number of species out of the 19 total which share a moderately high abundance as indicated by the curve patterns observed in the Figure (Fig. 4.7.1).
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Table 4.7.1 Rank abundance curves of species by site.
Rank VQS PIK GLA JSM
1 Umbilicaria virginis Umbilicaria virginis Umbilicaria virginis Acarospora contigua
2 Xanthoria elegans Rhizocarpon geographicum Rhizocarpon geographicum Lecidea atrobrunnea
3 Lecidea atrobrunnea Lecidea atrobrunnea Lecidea atrobrunnea Xanthoria elegans
4 Aspicilia caesiocinerea Acarospora contigua Umbilicaria decussata Aspicilia caesiocinerea
5 Umbilicaria decussata Xanthoria elegans Acarospora contigua Rhizocarpon geographicum
6 Acarospora contigua Lecanora rupicola Lecidea tessellata Umbilicaria virginis
7 Lecanora rupicola Aspicilia caesiocinerea Lecanora rupicola Xanthoparmelia cumberlandia
8 Xanthoparmelia wyomingica Umbilicaria decussata Aspicilia caesiocinerea Lecanora rupicola
9 Rhizocarpon geographicum Lecidea tessellata Xanthoria elegans Rhizoplaca chrysoleuca
10 Lecidea tessellata Rhizoplaca chrysoleuca Rhizoplaca chrysoleuca Xanthoparmelia wyomingica
11 Xanthoparmelia cumberlandia Pseudephebe minuscula Umbilicaria americana Xanthoparmelia chlorochroa
12 Rhizoplaca chrysoleuca Xanthoparmelia wyomingica Pseudephebe minuscula Umbilicaria americana
13 Xanthoparmelia chlorochroa Xanthoparmelia cumberlandia Xanthoparmelia wyomingica Pseudephebe minuscula
14 Pseudephebe minuscula Diploschistes muscorum Xanthoparmelia cumberlandia Lecidea tessellata
15 Diploschistes muscorum Xanthoparmelia chlorochroa Xanthoparmelia chlorochroa Umbilicaria decussata
16 Vulpicida tilesii Umbilicaria americana Diploschistes muscorum Diploschistes muscorum
17 Umbilicaria americana Vulpicida tilesii Thamnolia subuliformis Thamnolia subuliformis
18 Thamnolia subuliformis Thamnolia subuliformis Vulpicida pinastri Vulpicida pinastri
19 Vulpicida pinastri (0) Vulpicida pinastri Vulpicida tilesii Vulpicida tilesii
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Rank Abundance Curves
Abundance Rank
Figure 4.7.1 Graph displaying the rank abundance curves of lichen species (1-19) (proportional abundance with respect to rank abundance) per site.
4,8 SPSS
Final statistical analysis for the site and aspect species comparisons was carried out in IBMs SPSS statistical software (see Tables 4.8.1 through 4.8.5). With species site totals ranging from 0 to 3575, these were not normal distributions and the raw count also exhibited overdispersion. Trying to transform count data does not reliably improve analyses, necessitating a more conservative approach to analysis (O'Hara and Kotze 2010). A
26


Generalized Linear Model (GLIM) is a very flexible statistical model that works well with distributed dependent variables and allows for customization within the software itself. It also allows accommodation for empty cells (zero count values) and for customization to refine outputs within the study. For this study, species count was the dependent variable to be assessed against the four sites and their respective aspects for a total case processing value of 570 (19 species by 30 (summed from upper/lower aspects from all fours sites)). Individual species counts contained a minimum of 0 and a maximum value of 1,037 (Table 4.7.1). A negative binomial probability distribution with log link function was assumed, due to the fact that this count data exhibited a highly non-normal distribution. All assumptions were under the null hypothesis that no significant difference in species counts from site to site and when comparing aspects would be found. This assumption was made because of the nature of the ecosystems contained in this study. As mentioned before, these are harsh environments, so diversity can be convoluted in comparison to lower elevation systems (Brodo et al. 2001b). The goodness of fit displays how well the values fit the distribution of the assumption of negative binomial with a log link (Table 4.7.3). Initially, pairwise comparisons were run putting each site up against the others. This was executed based on the dependent variable of count (species) first against site comparisons then against site aspects (Table 4.8.6). The Bonferroni correction was chosen as a more conservative approach over the least significant difference (LSD) model as a correction for multiple comparisons. This model, however, was not deemed appropriate because it resulted in everything showing significance. The pairwise comparisons were then run with the Bonferroni correction, which corrects for post-hoc multiple comparisons. Using the Bonferroni correction for pairwise comparisons of differences in counts between sites or aspects, values below the p= 0.05 level were
27


considered significant. With the site pairwise comparisons, only the sites PIK and JSM, when compared, were not statistically different (p=0.736). The total counts at these two sites were 13,160 and 11,800, respectively. The rest of the comparisons all showed significant differences in lichen abundance between sites (Table 4.8.6). Finally, pairwise comparisons were conducted between aspects, and only the South to East comparisons showed any significance.
Table 4.8.1 This table display the overall data count with its minimum, maximum, mean, and standard deviation
Continuous Variable Information
N Minimum Maximum Mean Std. Deviation
Dependent Count 570 Variable 0 1037 88.43 138.982
Table 4.8.2 Variables per site. Note GLA has less due to the East aspect being excluded (Site Quadrats x 19 Species) Categorical Variable Information
N
Factor Site VQS 152
PIK 152
GLA 114
JSM 152
Total 570
Aspect West 152
South 152
North 152
East 114
Total 570
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Table 4.8.3 This table displays how well the data values fit the distribution with dependent variable count modeled against site and aspect Goodness of Fit8
Value df Value/df
Deviance 2011.923 563 3.574
Scaled Deviance 2011.923 563
Pearson Chi-Square 1200.764 563 2.133
Scaled Pearson Chi-Square 1200.764 563
Log Likelihoodb -3082.607
Dependent Variable: Count Model: (Intercept), Site, Aspect3
a. Information criteria are in smaller-is-better form.
b. The full log likelihood function is displayed and used in computing information criteria.
Table 4.8.4 Omnibus testing fitted model against intercept-only models (site to aspect). Model effects tests the intercept model against site and aspect using negative binomial regression
Omnibus Test8 Tests of Model Effects
Likelihood Ratio Chi-Square df Sig. Source Type III
90.971 6 0.000 Wald Chi-Square df Sig.
Dependent Variable: Count Model: (Intercept), Site, Aspect3 (Intercept) 10509.880 1 0.000
a. Compares the fitted model against the intercept-only model. Site 62.927 3 0.000
Aspect 16.959 3 0.001
Dependent Variable: Count Model: (Intercept), Site, Aspect
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Table 4.8.5 the reported mean and standard error for each site along with the 95% Wald confidence interval for upper and lower quadrats
Estimated Marginal Means Site/Aspect
Site Mean Std. Error 95% Wald Confidence Interval
Lower Upper
VQS 51.34 4.220 43.70 60.32
PIK 89.82 7.475 76.30 105.73
GLA 135.78 13.338 112.00 164.61
JSM 74.65 6.183 63.46 87.80
Aspect Mean Std. Error 95% Wald Confidence Interval
Lower Upper
West 90.57 7.417 77.14 106.34
South 104.08 8.504 88.68 122.15
North 80.39 6.616 68.41 94.46
East 61.68 6.223 50.61 75.17
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Table 4.8.6 Site to site comparisons using species count data as the dependent variable and the Bonferroni correction for multiple comparisons
Pairwise Comparisons Site
(I) Site Mean Difference (I-J) Std. Error df Bonferroni Sig. 95% Wald Confidence Interval for Difference
Lower Upper
VQS PIK -38.47a 8.629 1 0.000 -61.24 -15.71
GLA -84.44a 13.911 1 0.000 -121.14 -47.74
JSM -23.30a 7.478 1 0.011 -43.03 -3.57
PIK VQS 38.47a 8.629 1 0.000 15.71 61.24
GLA -45.97a 15.657 1 0.020 -87.27 -4.66
JSM 15.17 9.826 1 0.736 -10.75 41.09
GLA VQS 84.44a 13.911 1 0.000 47.74 121.14
PIK 45.97a 15.657 1 0.020 4.66 87.27
JSM 61.14a 14.494 1 0.000 22.90 99.38
JSM VQS 23.30a 7.478 1 0.011 3.57 43.03
PIK -15.17 9.826 1 0.736 -41.09 10.75
GLA -61.14a 14.494 1 0.000 -99.38 -22.90
Pairwise comparisons of estimated marginal means based on the original scale of dependent variable Count a. The mean difference is significant at the .05 level.
Pairwise Comparisons Aspect
(I) Aspect Mean Difference (I-J) Std. Error df Bonferroni Sig. 95% Wald Confidence Interval for Difference
Lower Upper
West South -13.50 11.255 1 1.000 -43.20 16.19
North 10.19 9.937 1 1.000 -16.03 36.40
East 28.89a 9.796 1 0.019 3.05 54.74
South West 13.50 11.255 1 1.000 -16.19 43.20
North 23.69 10.784 1 0.168 -4.76 52.14
East 42.40a 10.611 1 0.000 14.40 70.39
North West -10.19 9.937 1 1.000 -36.40 16.03
South -23.69 10.784 1 0.168 -52.14 4.76
East 18.71 9.103 1 0.239 -5.31 42.72
East West -28.89a 9.796 1 0.019 -54.74 -3.05
South -42.40a 10.611 1 0.000 -70.39 -14.40
North -18.71 9.103 1 0.239 -42.72 5.31
Pairwise comparisons of estimated marginal means based on the original scale of dependent variable Count
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CHAPTER V
DISCUSSION
This study originally sought out to discover relationships of biological soil crusts (BSCs) in relation to high elevation sites in regards to (e.g., alpine and subalpine biomes). Once in situ, however, it was quickly discovered that the ground cover in these biomes contained few (if any) BSCs, but very abundant with lichen cover. Upon discovering this, the research switched tracks, focusing on lichen exclusively. Within the sites themselves, the geology and land cover was fairly homogenous, especially in comparing the VQS and PIK site, with the only difference being PIK hosting a larger congregation of metamorphic rocks. Overall, the geology remained similar, containing granite, gneiss, schist, and quartz outcrops.
The subalpine site held the least amount of rock cover for lichen habitability, and more particularly on the eastern side of JSM. This aspect hosted small areas of grass, but highly abundant soil patches, as if some process, such as a large herd of mammals, had disturbed the area. This aspect also contained the lowest count (77). No other aspect at any of the other sites had such drastically low lichen counts. The three alpine sites (VQS, PIK,
GLA) ran along a ridge from North to South. Both VQS and GLA had their highest species counts on southerly aspects, with GLA containing the largest overall site total (17,303).
When comparing the land cover between sites, GLA also had the most surface area of rocks than any other study site. The majority of lichen species at the GLA site were also known for colonizing rock faces. This is typical of above tree line ecosystems where mostly rocky ground remains the main landscape feature. Both Vulpicidapinastri/tilesii and Thamnolia subuliformis are well known soil-based species, and Xanthoparmeliawyomingica is known to colonize both soil and rock. The high species count at GLA is most likely a result of these
32


two environmental factors (i.e., greater rock surface areas and the ability of certain species to thrive on rock specifically). The overall lichen species count totals seemed to lend more to differences among sites than some of the tests ran. For example, in regards to the Simpsons Diversity Index, the overall diversity yields variance across sites basically an 87% chance that any two random species chosen would have similar presence. Jaccards showed that there was high species overlap across all four sites, which is to be expected on a smaller areal study with only 19 species of lichen identified. There was, however, a large overlap when comparing the east and south aspects among the means of alpine to subalpine. This was prevalent in the pairwise aspect comparisons which were significant using the Bonferroni adjustment scores. Especially interesting, this analysis seemed to follow the higher totals for east and south aspects. In reviewing historical Google Earth imagery of the sites, a recurring snow bank was noticed, sometimes extending post winter along the eastern aspect of the ridgeline. The western aspect tends to experience a westerly wind pattern typical for that area of the Rocky Mountains, and this phenomenon, coupled with a cleared west side and higher snow accumulation on the east side, may be what could be affecting lichen totals. To determine if this is indeed a contributing factor to lichen numbers and colonization, further (snowfall) studies need to be conducted.
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CHAPTER VI
CONCLUSION
High alpine ecosystems, such as those studied for this research, represent harsh environments for any organisms. Exposure in these locales remains a constant: from desiccating winds and UV rays, to drastic temperature fluctuations-, all inescapable on the bare earth with no shade. Resilience through biological adaptation represents a key component for species survival in such harsh extremes. Lichen, with their symbiotic relationship of algae/cyanobacteria housed inside a fungal body have managed to survive in all ecosystems across all continents on this planet. At its core, this study set out to discover spatial characteristics of such species. Though lichen dominated the area of study in between ground level vascular plants, this study teased out a few relational findings among the 19 predominant species identified at the four study sites.
One analysis that was briefly looked at, but not imposed, was slope analysis. ESRIs Arcmaps slope analysis tool, when applied to the study areas, showed all sites fell into a similar percentage (17-22%) across sites. The only differences were at VQS, which had (0-7%), the West side of JSM (22-26%) and the east aspect of GLA (31-37%), which was too steep to assess without the aid of climbing gear. These slopes were not steep enough to create a rock fall situation that may have affected the overall rock colonization of lichen present. In fact, most rocks were embedded in the ground at these sites. Due to the general evenness of slope across all four sites, then, slope analyses were left out of this study. If proper safety gear could be utilized for assessing the east aspect of GLA, however, then slope could be worth investigating in a future follow-up study. This could enhance findings in relation to the aspects and their influence on colonization.
34


As the south and east facing aspects contained more total lichen than most of the others, other factors could be included in future studies, as a wide range of varying climatic factors (Grabherr, Gottfried and Pauli 2000) would likely yield aspect relationships. For example, surrounding vegetation helps regulate moisture and soil temperature (Aalto, le Roux and Luoto 2013), and exposure to the sun and its path across the sky, will likely have an effect on those elements. Typical westerly winds for the Rocky Mountain range would also likely play a part, as noted in the discussion and Google Earth images in Appendices F, when northern aspects remain cooler and moister than warmer southerly exposures (Selvakumar et al. 2009). Those climatic factors, in conjunction with, lichen habitat preferences, could infer aspect-based populations based on solar and climatic data in relation to each of the four sites assessed.
In addition to slope and aspect for possible future research, the effects of carbon sequestration and nitrogen fixation could also be investigated. Most studies of this nature are conducted on soil-based species only (De Deyn, Cornelissen and Bardgett 2008), and most often in controlled laboratory environments (Yoshitake et al. 2010, Barger et al. 2006) where inputs into the soil can be measured. Inputs and variables associated with this aspect remain diverse: light, temperature, radiation, moisture, type of cyanobacteria or green algae, morphology, snow cover, and permafrost, among others (Lange 2002, Lange and Green 1996, De Deyn et al. 2008). These prove difficult to measure due to the abundance of external influences (Lange and Green 2005, Lange, Reichenberger and Walz 1997). Though this study was not geared for such assessments, each of the above-mentioned components could represent important topics for future research endeavors.
35


In the end, while this study laid the basic foundation for future research endeavors related to alpine and sub-alpine lichen spatiality and identification, it also tested several statistical measures against somewhat large spatial parameters. This will allow future alpine and sub-alpine lichen research to focus on more specific correlations, including climate-related phenomena. Overall, this study will also inform other and further studies on lichen in alpine and sub-alpine ecosystems of Rocky Mountain National Park specifically, and North America and other continents more generally.
36


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42


APPENDIX
A. LICHEN LIST FROM RMNP
https://www.nps.gov/romo/learn/nature/looking_for_lichens.htm
Scientific Name Common Name
Acarosporct molybdina
Acarosporct schleicheri
Allocetraria madreporiformis
Arctoparmelia subcentrifiiga
Arthonia sp.
As pi cilia caesiocinerea
Aspicilia calcarea
Aspicilia sublapponica
Bacidia subincompta
Brodoa oroarctica
Bryoria fusee scens Pale-footed horsehair lichen
Buellia leptocline
Caloplaca exsecuta
Caloplaca sinapisperma
Caloplaca sp. 1
Caloplaca sp. 2
Caloplaca sp. 3
Caloplaca sp. 4
Caloplaca vitellinula
Calvitimela armeniaca
Candelariella sp.
Candelariella terrigena
Carbonea vorticosa
Catapyrenium daedaleum
Cetraria aculeata
Cetraria ericetorum Iceland lichen
Cetraria islandica
Chaenotheca ferruginea
Chromatochlamys muscorum octospora
Cladonia cariosa
Cladonia cenotea Powdered funnel lichen
Cladonia cervicomis Ladder lichen
Cladonia chlorophaea group
Cladonia coccifera
43


Cladonia coniocraea Common powderhom
Cladonia comuta Bighorn lichen
Cladonia comuta groenlandica
Cladonia deformis or sulphurina
Cladonia didyma Southern soldiers
Cladonia ecmocyna Frosted cladonia
Cladonia ecmocyna intermedia
Cladonia gracilis
Cladonia macilenta bacillaris
Cladonia macroceras
Cladonia macrophyllodes Farge-leaved cladonia
Cladonia phyllophora
Cladonia sp. 1
Cladonia sp. 2
Cladonia sp. 3
Cladonia sp. 4
Cladonia sulphurina
Cypehlium notarisii
Dermatocarpon miniatum Feather lichen
Dermatocarpon reticulatum
Diploschistes muscorum Cowpie lichen
Evemia divaricata Mountain oakmoss lichen
Flavocetraria cucullata
Flavocetraria nivalis Crinkled snow lichen
Flavoparmelia caperata
Flavoparmelia mtidota
Hypogymnia bitteri
Hypogymnia farinacea
Icmadophila ericetomm
Imshaugia aleurites
Ionaspis lacustris
Fasallia papulosa
Fasallia pensylvanica
Leccmora allophana
Leccmora chloropolia
Leccmora epibryon
Leccmora impudens
Leccmora novomexicana
Leccmora polytropa
Leccmora mpicola
Leccmora sp.
44


Leccmora thomsonii
Lecidect atrobrunnea
Lecidect promiscens
Lecidect sp.
Lecidect tessellata Tile lichen
Lecidella scabra
Leciophysma furfurascens
Lepraria sp.
Melanelia exasperatula Lustrous camouflage lichen
Melanelia glabroides
Melanelia subolivacea Brown-eyed camouflage lichen
Micarea tuberculata
Miriquidica garovaglii
Mycobilimbia berengeriana
Nephroma bellum
Nephroma parile Powdery kidney lichen
Ochrolechia upsaliensis
Parmelia fraudans
Parmelia saxatilis Salted shield lichen
Parmelia sulcata Hammered shield lichen
Parmeliopsis ambigua Green starburst lichen
Parmeliopsis hyperopta Gray starburst lichen
Peltigera aphthosa Felt lichen
Peltigera canina Dog lichen
Peltigera collina
Peltigera horizontalis polydactyla
Peltigera malacea Veinless pelt
Peltigera rufescens Field dog lichen
Peltigera sp.
Peltigera venosa
Phaeophyscia constipata
Physcia stellaris Star rosette lichen
Physconia muscigena
Placidium lachneum
Placynthium nigrum
Polyblastia cucurbitula
Polychidium muscicola
Porocyphus dispersus
Porpidia superba
Porpidia thomsonii
Protoparmelia atriseda
45


Protoparmelia badia
Pseudephebe minuscula
Pseudephebe pubescens?
Psora luridella
Psora montana
Psoroma hypnorum Green moss-shingle
Punctelia perreticulata
Rhizocctrpon disporum
Rhizocctrpon geographicum Yellow map lichen
Rhizoplaca chrysoleuca
Rhizoplaca melanophthalma
Rinodina mniaraea
Rinodina turfacea Tundra pepper-spore lichen
Schaereria cinereorufa
Solorina crocea Orange chocolate chip lichen
Staurothele clopimoides
Stereocaulon sp.
Stereocaulon tomentosum group
Thamnolia subuliformis
Toninia tristis
Trapeliopsis granulosa Mottled-disk lichen
Umbilicaria americana Frosted rock tripe
Umbilicaria decussata
Umbilicaria deusta
Umbilicaria hyperborea Blistered rock tripe
Umbilicaria krascheninnikovii Salty rock tripe
Umbilicaria lyngei
Umbilicaria torrefacta
Umbilicaria vellea Alpine rock tripe
Umbilicaria virginis
Usnea cavernosa
Usnea lapponica Powdered beard lichen
Vulpicida pinastri Powdered sunshine lichen
Vulpicida tilesii
Xanthoparmelia coloradoensis Colorado rock-shield lichen
Xanthoparmelia coloradoensis or cumberlandia
Xanthoparmelia sp.
Xanthoparmelia wyomingica Shingled rock-shield lichen
Xanthoria sorediata
46


B. SPECIES TABLES
Morphology Species Site Aspect Elevation Count
Crustose Acarospora contigua VQS West Upper 76
Crustose Acarospora contigua VQS West Lower 95
Crustose Acarospora contigua VQS South Upper 57
Crustose Acarospora contigua VQS South Lower 120
Crustose Acarospora contigua VQS North Upper 60
Crustose Acarospora contigua VQS North Lower 91
Crustose Acarospora contigua VQS East Upper 65
Crustose Acarospora contigua VQS East Lower 57

Crustose Acarospora contigua PIK West Upper 97
Crustose Acarospora contigua PIK West Lower 101
Crustose Acarospora contigua PIK South Upper 127
Crustose Acarospora contigua PIK South Lower 52
Crustose Acarospora contigua PIK North Upper 85
Crustose Acarospora contigua PIK North Lower 141
Crustose Acarospora contigua PIK East Upper 278
Crustose Acarospora contigua PIK East Lower 275

Crustose Acarospora contigua GLA West Upper 131
Crustose Acarospora contigua GLA West Lower 198
Crustose Acarospora contigua GLA South Upper 201
Crustose Acarospora contigua GLA South Lower 236
Crustose Acarospora contigua GLA North Upper 157
Crustose Acarospora contigua GLA North Lower 210
Crustose Acarospora contigua GLA East Upper N/A
Crustose Acarospora contigua GLA East Lower N/A

Crustose Acarospora contigua JSM West Upper 258
Crustose Acarospora contigua JSM West Lower 323
Crustose Acarospora contigua JSM South Upper 407
Crustose Acarospora contigua JSM South Lower 439
Crustose Acarospora contigua JSM North Upper 496
Crustose Acarospora contigua JSM North Lower 465
Crustose Acarospora contigua JSM East Upper 317
Crustose Acarospora contigua JSM East Lower 29
47


Morphology Species Site Aspect Elevation Count
Crustose Aspicilia caesiocinerea VQS West Upper 131
Crustose Aspicilia caesiocinerea VQS West Lower 75
Crustose Aspicilia caesiocinerea VQS South Upper 208
Crustose Aspicilia caesiocinerea VQS South Lower 140
Crustose Aspicilia caesiocinerea VQS North Upper 88
Crustose Aspicilia caesiocinerea VQS North Lower 57
Crustose Aspicilia caesiocinerea VQS East Upper 146
Crustose Aspicilia caesiocinerea VQS East Lower 88

Crustose Aspicilia caesiocinerea PIK West Upper 15
Crustose Aspicilia caesiocinerea PIK West Lower 9
Crustose Aspicilia caesiocinerea PIK South Upper 220
Crustose Aspicilia caesiocinerea PIK South Lower 29
Crustose Aspicilia caesiocinerea PIK North Upper 168
Crustose Aspicilia caesiocinerea PIK North Lower 182
Crustose Aspicilia caesiocinerea PIK East Upper 227
Crustose Aspicilia caesiocinerea PIK East Lower 46

Crustose Aspicilia caesiocinerea GLA West Upper 42
Crustose Aspicilia caesiocinerea GLA West Lower 122
Crustose Aspicilia caesiocinerea GLA South Upper 140
Crustose Aspicilia caesiocinerea GLA South Lower 137
Crustose Aspicilia caesiocinerea GLA North Upper 44
Crustose Aspicilia caesiocinerea GLA North Lower 258
Crustose Aspicilia caesiocinerea GLA East Upper N/A
Crustose Aspicilia caesiocinerea GLA East Lower N/A

Crustose Aspicilia caesiocinerea JSM West Upper 226
Crustose Aspicilia caesiocinerea JSM West Lower 205
Crustose Aspicilia caesiocinerea JSM South Upper 350
Crustose Aspicilia caesiocinerea JSM South Lower 103
Crustose Aspicilia caesiocinerea JSM North Upper 155
Crustose Aspicilia caesiocinerea JSM North Lower 201
Crustose Aspicilia caesiocinerea JSM East Upper 126
Crustose Aspicilia caesiocinerea JSM East Lower 2
48


Morphology Species Site Aspect Elevation Count
Crustose Diploschistes muscorum VQS West Upper 0
Crustose Diploschistes muscorum VQS West Lower 0
Crustose Diploschistes muscorum VQS South Upper 3
Crustose Diploschistes muscorum VQS South Lower 17
Crustose Diploschistes muscorum VQS North Upper 0
Crustose Diploschistes muscorum VQS North Lower 0
Crustose Diploschistes muscorum VQS East Upper 0
Crustose Diploschistes muscorum VQS East Lower 0

Crustose Diploschistes muscorum PIK West Upper 0
Crustose Diploschistes muscorum PIK West Lower 22
Crustose Diploschistes muscorum PIK South Upper 0
Crustose Diploschistes muscorum PIK South Lower 6
Crustose Diploschistes muscorum PIK North Upper 0
Crustose Diploschistes muscorum PIK North Lower 0
Crustose Diploschistes muscorum PIK East Upper 0
Crustose Diploschistes muscorum PIK East Lower 1

Crustose Diploschistes muscorum GLA West Upper 1
Crustose Diploschistes muscorum GLA West Lower 17
Crustose Diploschistes muscorum GLA South Upper 3
Crustose Diploschistes muscorum GLA South Lower 1
Crustose Diploschistes muscorum GLA North Upper 1
Crustose Diploschistes muscorum GLA North Lower 4
Crustose Diploschistes muscorum GLA East Upper N/A
Diploschistes muscorum GLA East Lower N/A

Crustose Diploschistes muscorum JSM West Upper 18
Crustose Diploschistes muscorum JSM West Lower 5
Crustose Diploschistes muscorum JSM South Upper 5
Crustose Diploschistes muscorum JSM South Lower 0
Crustose Diploschistes muscorum JSM North Upper 9
Crustose Diploschistes muscorum JSM North Lower 0
Crustose Diploschistes muscorum JSM East Upper 0
Crustose Diploschistes muscorum JSM East Lower 0
49


Morphology Species Site Aspect Elevation Count
Crustose Lecanora rupicola VQS West Upper 23
Crustose Lecanora rupicola VQS West Lower 32
Crustose Lecanora rupicola VQS South Upper 42
Crustose Lecanora rupicola VQS South Lower 26
Crustose Lecanora rupicola VQS North Upper 44
Crustose Lecanora rupicola VQS North Lower 56
Crustose Lecanora rupicola VQS East Upper 29
Crustose Lecanora rupicola VQS East Lower 30

Crustose Lecanora rupicola PIK West Upper 157
Crustose Lecanora rupicola PIK West Lower 91
Crustose Lecanora rupicola PIK South Upper 125
Crustose Lecanora rupicola PIK South Lower 61
Crustose Lecanora rupicola PIK North Upper 93
Crustose Lecanora rupicola PIK North Lower 84
Crustose Lecanora rupicola PIK East Upper 232
Crustose Lecanora rupicola PIK East Lower 113

Crustose Lecanora rupicola GLA West Upper 140
Crustose Lecanora rupicola GLA West Lower 185
Crustose Lecanora rupicola GLA South Upper 246
Crustose Lecanora rupicola GLA South Lower 144
Crustose Lecanora rupicola GLA North Upper 118
Crustose Lecanora rupicola GLA North Lower 133
Crustose Lecanora rupicola GLA East Upper N/A
Crustose Lecanora rupicola GLA East Lower N/A

Crustose Lecanora rupicola JSM West Upper 70
Crustose Lecanora rupicola JSM West Lower 62
Crustose Lecanora rupicola JSM South Upper 71
Crustose Lecanora rupicola JSM South Lower 10
Crustose Lecanora rupicola JSM North Upper 127
Crustose Lecanora rupicola JSM North Lower 139
Crustose Lecanora rupicola JSM East Upper 47
Crustose Lecanora rupicola JSM East Lower 5
50


Morphology Species Site Aspect Elevation Count
Crustose Lecidea atrobrunnea VQS West Upper 119
Crustose Lecidea atrobrunnea VQS West Lower 87
Crustose Lecidea atrobrunnea VQS South Upper 177
Crustose Lecidea atrobrunnea VQS South Lower 190
Crustose Lecidea atrobrunnea VQS North Upper 191
Crustose Lecidea atrobrunnea VQS North Lower 104
Crustose Lecidea atrobrunnea VQS East Upper 140
Crustose Lecidea atrobrunnea VQS East Lower 98

Crustose Lecidea atrobrunnea PIK West Upper 151
Crustose Lecidea atrobrunnea PIK West Lower 191
Crustose Lecidea atrobrunnea PIK South Upper 314
Crustose Lecidea atrobrunnea PIK South Lower 124
Crustose Lecidea atrobrunnea PIK North Upper 150
Crustose Lecidea atrobrunnea PIK North Lower 151
Crustose Lecidea atrobrunnea PIK East Upper 356
Crustose Lecidea atrobrunnea PIK East Lower 178

Crustose Lecidea atrobrunnea GLA West Upper 489
Crustose Lecidea atrobrunnea GLA West Lower 505
Crustose Lecidea atrobrunnea GLA South Upper 627
Crustose Lecidea atrobrunnea GLA South Lower 517
Crustose Lecidea atrobrunnea GLA North Upper 349
Crustose Lecidea atrobrunnea GLA North Lower 411
Crustose Lecidea atrobrunnea GLA East Upper N/A
Crustose Lecidea atrobrunnea GLA East Lower N/A

Crustose Lecidea atrobrunnea JSM West Upper 235
Crustose Lecidea atrobrunnea JSM West Lower 179
Crustose Lecidea atrobrunnea JSM South Upper 256
Crustose Lecidea atrobrunnea JSM South Lower 279
Crustose Lecidea atrobrunnea JSM North Upper 285
Crustose Lecidea atrobrunnea JSM North Lower 352
Crustose Lecidea atrobrunnea JSM East Upper 189
Crustose Lecidea atrobrunnea JSM East Lower 1
51


Morphology Species Site Aspect Elevation Count
Crustose Lecidea tessellata VQS West Upper 20
Crustose Lecidea tessellata VQS West Lower 35
Crustose Lecidea tessellata VQS South Upper 42
Crustose Lecidea tessellata VQS South Lower 44
Crustose Lecidea tessellata VQS North Upper 34
Crustose Lecidea tessellata VQS North Lower 17
Crustose Lecidea tessellata VQS East Upper 38
Crustose Lecidea tessellata VQS East Lower 7

Crustose Lecidea tessellata PIK West Upper 29
Crustose Lecidea tessellata PIK West Lower 37
Crustose Lecidea tessellata PIK South Upper 94
Crustose Lecidea tessellata PIK South Lower 53
Crustose Lecidea tessellata PIK North Upper 93
Crustose Lecidea tessellata PIK North Lower 110
Crustose Lecidea tessellata PIK East Upper 133
Crustose Lecidea tessellata PIK East Lower 87

Crustose Lecidea tessellata GLA West Upper 102
Crustose Lecidea tessellata GLA West Lower 187
Crustose Lecidea tessellata GLA South Upper 185
Crustose Lecidea tessellata GLA South Lower 162
Crustose Lecidea tessellata GLA North Upper 126
Crustose Lecidea tessellata GLA North Lower 223
Crustose Lecidea tessellata GLA East Upper N/A
Crustose Lecidea tessellata GLA East Lower N/A

Crustose Lecidea tessellata JSM West Upper 0
Crustose Lecidea tessellata JSM West Lower 0
Crustose Lecidea tessellata JSM South Upper 0
Crustose Lecidea tessellata JSM South Lower 0
Crustose Lecidea tessellata JSM North Upper 50
Crustose Lecidea tessellata JSM North Lower 0
Crustose Lecidea tessellata JSM East Upper 35
Crustose Lecidea tessellata JSM East Lower 6
52


Morphology Species Site Aspect Elevation Count
Fruticose Pseudephebe minuscula VQS West Upper 4
Fruticose Pseudephebe minuscula VQS West Lower 4
Fruticose Pseudephebe minuscula VQS South Upper 0
Fruticose Pseudephebe minuscula VQS South Lower 6
Fruticose Pseudephebe minuscula VQS North Upper 5
Fruticose Pseudephebe minuscula VQS North Lower 1
Fruticose Pseudephebe minuscula VQS East Upper 0
Fruticose Pseudephebe minuscula VQS East Lower 0

Fruticose Pseudephebe minuscula PIK West Upper 23
Fruticose Pseudephebe minuscula PIK West Lower 22
Fruticose Pseudephebe minuscula PIK South Upper 78
Fruticose Pseudephebe minuscula PIK South Lower 11
Fruticose Pseudephebe minuscula PIK North Upper 6
Fruticose Pseudephebe minuscula PIK North Lower 24
Fruticose Pseudephebe minuscula PIK East Upper 82
Fruticose Pseudephebe minuscula PIK East Lower 0

Fruticose Pseudephebe minuscula GLA West Upper 40
Fruticose Pseudephebe minuscula GLA West Lower 18
Fruticose Pseudephebe minuscula GLA South Upper 50
Fruticose Pseudephebe minuscula GLA South Lower 37
Fruticose Pseudephebe minuscula GLA North Upper 5
Fruticose Pseudephebe minuscula GLA North Lower 12
Fruticose Pseudephebe minuscula GLA East Upper N/A
Fruticose Pseudephebe minuscula GLA East Lower N/A

Fruticose Pseudephebe minuscula JSM West Upper 35
Fruticose Pseudephebe minuscula JSM West Lower 3
Fruticose Pseudephebe minuscula JSM South Upper 7
Fruticose Pseudephebe minuscula JSM South Lower 0
Fruticose Pseudephebe minuscula JSM North Upper 0
Fruticose Pseudephebe minuscula JSM North Lower 5
Fruticose Pseudephebe minuscula JSM East Upper 42
Fruticose Pseudephebe minuscula JSM East Lower 0
53


Morphology Species Site Aspect Elevation Count
Crustose Rhizocarpon geographicum VQS West Upper 63
Crustose Rhizocarpon geographicum VQS West Lower 3
Crustose Rhizocarpon geographicum VQS South Upper 28
Crustose Rhizocarpon geographicum VQS South Lower 27
Crustose Rhizocarpon geographicum VQS North Upper 63
Crustose Rhizocarpon geographicum VQS North Lower 27
Crustose Rhizocarpon geographicum VQS East Upper 30
Crustose Rhizocarpon geographicum VQS East Lower 7

Crustose Rhizocarpon geographicum PIK West Upper 351
Crustose Rhizocarpon geographicum PIK West Lower 222
Crustose Rhizocarpon geographicum PIK South Upper 450
Crustose Rhizocarpon geographicum PIK South Lower 48
Crustose Rhizocarpon geographicum PIK North Upper 124
Crustose Rhizocarpon geographicum PIK North Lower 201
Crustose Rhizocarpon geographicum PIK East Upper 648
Crustose Rhizocarpon geographicum PIK East Lower 316

Crustose Rhizocarpon geographicum GLA West Upper 718
Crustose Rhizocarpon geographicum GLA West Lower 579
Crustose Rhizocarpon geographicum GLA South Upper 869
Crustose Rhizocarpon geographicum GLA South Lower 633
Crustose Rhizocarpon geographicum GLA North Upper 307
Crustose Rhizocarpon geographicum GLA North Lower 398
Crustose Rhizocarpon geographicum GLA East Upper N/A
Crustose Rhizocarpon geographicum GLA East Lower N/A

Crustose Rhizocarpon geographicum JSM West Upper 125
Crustose Rhizocarpon geographicum JSM West Lower 118
Crustose Rhizocarpon geographicum JSM South Upper 185
Crustose Rhizocarpon geographicum JSM South Lower 5
Crustose Rhizocarpon geographicum JSM North Upper 174
Crustose Rhizocarpon geographicum JSM North Lower 426
Crustose Rhizocarpon geographicum JSM East Upper 232
Crustose Rhizocarpon geographicum JSM East Lower 31
54


Morphology Species Site Aspect Elevation Count
Foliose Rhizoplaca chrysoleuca VQS West Upper 4
Foliose Rhizoplaca chrysoleuca VQS West Lower 4
Foliose Rhizoplaca chrysoleuca VQS South Upper 3
Foliose Rhizoplaca chrysoleuca VQS South Lower 31
Foliose Rhizoplaca chrysoleuca VQS North Upper 41
Foliose Rhizoplaca chrysoleuca VQS North Lower 9
Foliose Rhizoplaca chrysoleuca VQS East Upper 23
Foliose Rhizoplaca chrysoleuca VQS East Lower 6

Foliose Rhizoplaca chrysoleuca PIK West Upper 87
Foliose Rhizoplaca chrysoleuca PIK West Lower 46
Foliose Rhizoplaca chrysoleuca PIK South Upper 63
Foliose Rhizoplaca chrysoleuca PIK South Lower 52
Foliose Rhizoplaca chrysoleuca PIK North Upper 50
Foliose Rhizoplaca chrysoleuca PIK North Lower 54
Foliose Rhizoplaca chrysoleuca PIK East Upper 63
Foliose Rhizoplaca chrysoleuca PIK East Lower 22

Foliose Rhizoplaca chrysoleuca GLA West Upper 55
Foliose Rhizoplaca chrysoleuca GLA West Lower 114
Foliose Rhizoplaca chrysoleuca GLA South Upper 133
Foliose Rhizoplaca chrysoleuca GLA South Lower 105
Foliose Rhizoplaca chrysoleuca GLA North Upper 50
Foliose Rhizoplaca chrysoleuca GLA North Lower 11
Foliose Rhizoplaca chrysoleuca GLA East Upper N/A
Foliose Rhizoplaca chrysoleuca GLA East Lower N/A

Foliose Rhizoplaca chrysoleuca JSM West Upper 47
Foliose Rhizoplaca chrysoleuca JSM West Lower 92
Foliose Rhizoplaca chrysoleuca JSM South Upper 10
Foliose Rhizoplaca chrysoleuca JSM South Lower 82
Foliose Rhizoplaca chrysoleuca JSM North Upper 49
Foliose Rhizoplaca chrysoleuca JSM North Lower 69
Foliose Rhizoplaca chrysoleuca JSM East Upper 0
Foliose Rhizoplaca chrysoleuca JSM East Lower 0
55


Morphology Species Site Aspect Elevation Count
Fruiticose Thamnolia subuliformis VQS West Upper 0
Fruiticose Thamnolia subuliformis VQS West Lower 0
Fruiticose Thamnolia subuliformis VQS South Upper 1
Fruiticose Thamnolia subuliformis VQS South Lower 0
Fruiticose Thamnolia subuliformis VQS North Upper 0
Fruiticose Thamnolia subuliformis VQS North Lower 0
Fruiticose Thamnolia subuliformis VQS East Upper 0
Fruiticose Thamnolia subuliformis VQS East Lower 0

Fruiticose Thamnolia subuliformis PIK West Upper 0
Fruiticose Thamnolia subuliformis PIK West Lower 3
Fruiticose Thamnolia subuliformis PIK South Upper 0
Fruiticose Thamnolia subuliformis PIK South Lower 2
Fruiticose Thamnolia subuliformis PIK North Upper 0
Fruiticose Thamnolia subuliformis PIK North Lower 0
Fruiticose Thamnolia subuliformis PIK East Upper 0
Fruiticose Thamnolia subuliformis PIK East Lower 0

Fruiticose Thamnolia subuliformis GLA West Upper 0
Fruiticose Thamnolia subuliformis GLA West Lower 0
Fruiticose Thamnolia subuliformis GLA South Upper 1
Fruiticose Thamnolia subuliformis GLA South Lower 0
Fruiticose Thamnolia subuliformis GLA North Upper 0
Fruiticose Thamnolia subuliformis GLA North Lower 0
Fruiticose Thamnolia subuliformis GLA East Upper N/A
Fruiticose Thamnolia subuliformis GLA East Lower N/A

Fruiticose Thamnolia subuliformis JSM West Upper 0
Fruiticose Thamnolia subuliformis JSM West Lower 0
Fruiticose Thamnolia subuliformis JSM South Upper 0
Fruiticose Thamnolia subuliformis JSM South Lower 0
Fruiticose Thamnolia subuliformis JSM North Upper 0
Fruiticose Thamnolia subuliformis JSM North Lower 0
Fruiticose Thamnolia subuliformis JSM East Upper 0
Fruiticose Thamnolia subuliformis JSM East Lower 0
56


Morphology Species Site Aspect Elevation Count
Foliose Umbilicaria americana VQS West Upper 0
Foliose Umbilicaria americana VQS West Lower 3
Foliose Umbilicaria americana VQS South Upper 0
Foliose Umbilicaria americana VQS South Lower 0
Foliose Umbilicaria americana VQS North Upper 0
Foliose Umbilicaria americana VQS North Lower 0
Foliose Umbilicaria americana VQS East Upper 0
Foliose Umbilicaria americana VQS East Lower 0

Foliose Umbilicaria americana PIK West Upper 11
Foliose Umbilicaria americana PIK West Lower 1
Foliose Umbilicaria americana PIK South Upper 3
Foliose Umbilicaria americana PIK South Lower 0
Foliose Umbilicaria americana PIK North Upper 0
Foliose Umbilicaria americana PIK North Lower 0
Foliose Umbilicaria americana PIK East Upper 5
Foliose Umbilicaria americana PIK East Lower 0

Foliose Umbilicaria americana GLA West Upper 12
Foliose Umbilicaria americana GLA West Lower 7
Foliose Umbilicaria americana GLA South Upper 71
Foliose Umbilicaria americana GLA South Lower 43
Foliose Umbilicaria americana GLA North Upper 32
Foliose Umbilicaria americana GLA North Lower 52
Foliose Umbilicaria americana GLA East Upper N/A
Foliose Umbilicaria americana GLA East Lower N/A

Foliose Umbilicaria americana JSM West Upper 47
Foliose Umbilicaria americana JSM West Lower 34
Foliose Umbilicaria americana JSM South Upper 8
Foliose Umbilicaria americana JSM South Lower 4
Foliose Umbilicaria americana JSM North Upper 9
Foliose Umbilicaria americana JSM North Lower 24
Foliose Umbilicaria americana JSM East Upper 0
Foliose Umbilicaria americana JSM East Lower 0
57


Morphology Species Site Aspect Elevation Count
Foliose Umbilicaria decussata VQS West Upper 197
Foliose Umbilicaria decussata VQS West Lower 63
Foliose Umbilicaria decussata VQS South Upper 169
Foliose Umbilicaria decussata VQS South Lower 202
Foliose Umbilicaria decussata VQS North Upper 108
Foliose Umbilicaria decussata VQS North Lower 21
Foliose Umbilicaria decussata VQS East Upper 3
Foliose Umbilicaria decussata VQS East Lower 8

Foliose Umbilicaria decussata PIK West Upper 115
Foliose Umbilicaria decussata PIK West Lower 82
Foliose Umbilicaria decussata PIK South Upper 185
Foliose Umbilicaria decussata PIK South Lower 24
Foliose Umbilicaria decussata PIK North Upper 48
Foliose Umbilicaria decussata PIK North Lower 100
Foliose Umbilicaria decussata PIK East Upper 132
Foliose Umbilicaria decussata PIK East Lower 1

Foliose Umbilicaria decussata GLA West Upper 347
Foliose Umbilicaria decussata GLA West Lower 327
Foliose Umbilicaria decussata GLA South Upper 410
Foliose Umbilicaria decussata GLA South Lower 76
Foliose Umbilicaria decussata GLA North Upper 371
Foliose Umbilicaria decussata GLA North Lower 150
Foliose Umbilicaria decussata GLA East Upper N/A
Umbilicaria decussata GLA East Lower N/A

Foliose Umbilicaria decussata JSM West Upper 24
Foliose Umbilicaria decussata JSM West Lower 5
Foliose Umbilicaria decussata JSM South Upper 0
Foliose Umbilicaria decussata JSM South Lower 5
Foliose Umbilicaria decussata JSM North Upper 0
Foliose Umbilicaria decussata JSM North Lower 17
Foliose Umbilicaria decussata JSM East Upper 0
Foliose Umbilicaria decussata JSM East Lower 0
58


Morphology Species Site Aspect Elevation Count
Foliose Umbilicaria virginis VQS West Upper 473
Foliose Umbilicaria virginis VQS West Lower 304
Foliose Umbilicaria virginis VQS South Upper 442
Foliose Umbilicaria virginis VQS South Lower 333
Foliose Umbilicaria virginis VQS North Upper 265
Foliose Umbilicaria virginis VQS North Lower 75
Foliose Umbilicaria virginis VQS East Upper 200
Foliose Umbilicaria virginis VQS East Lower 40

Foliose Umbilicaria virginis PIK West Upper 309
Foliose Umbilicaria virginis PIK West Lower 256
Foliose Umbilicaria virginis PIK South Upper 514
Foliose Umbilicaria virginis PIK South Lower 337
Foliose Umbilicaria virginis PIK North Upper 326
Foliose Umbilicaria virginis PIK North Lower 240
Foliose Umbilicaria virginis PIK East Upper 512
Foliose Umbilicaria virginis PIK East Lower 28

Foliose Umbilicaria virginis GLA West Upper 643
Foliose Umbilicaria virginis GLA West Lower 865
Foliose Umbilicaria virginis GLA South Upper 1037
Foliose Umbilicaria virginis GLA South Lower 436
Foliose Umbilicaria virginis GLA North Upper 225
Foliose Umbilicaria virginis GLA North Lower 369
Foliose Umbilicaria virginis GLA East Upper N/A
Foliose Umbilicaria virginis GLA East Lower N/A

Foliose Umbilicaria virginis JSM West Upper 283
Foliose Umbilicaria virginis JSM West Lower 184
Foliose Umbilicaria virginis JSM South Upper 145
Foliose Umbilicaria virginis JSM South Lower 81
Foliose Umbilicaria virginis JSM North Upper 57
Foliose Umbilicaria virginis JSM North Lower 78
Foliose Umbilicaria virginis JSM East Upper 0
Foliose Umbilicaria virginis JSM East Lower 0
59


Morphology Species Site Aspect Elevation Count
Foliose Vulpicida pinastri VQS West Upper 0
Foliose Vulpicida pinastri VQS West Lower 0
Foliose Vulpicida pinastri VQS South Upper 0
Foliose Vulpicida pinastri VQS South Lower 0
Foliose Vulpicida pinastri VQS North Upper 0
Foliose Vulpicida pinastri VQS North Lower 0
Foliose Vulpicida pinastri VQS East Upper 0
Foliose Vulpicida pinastri VQS East Lower 0

Foliose Vulpicida pinastri PIK West Upper 0
Foliose Vulpicida pinastri PIK West Lower 1
Foliose Vulpicida pinastri PIK South Upper 3
Foliose Vulpicida pinastri PIK South Lower 0
Foliose Vulpicida pinastri PIK North Upper 0
Foliose Vulpicida pinastri PIK North Lower 0
Foliose Vulpicida pinastri PIK East Upper 0
Foliose Vulpicida pinastri PIK East Lower 0

Foliose Vulpicida pinastri GLA West Upper 0
Foliose Vulpicida pinastri GLA West Lower 0
Foliose Vulpicida pinastri GLA South Upper 0
Foliose Vulpicida pinastri GLA South Lower 0
Foliose Vulpicida pinastri GLA North Upper 0
Foliose Vulpicida pinastri GLA North Lower 0
Foliose Vulpicida pinastri GLA East Upper N/A
Foliose Vulpicida pinastri GLA East Lower N/A

Foliose Vulpicida pinastri JSM West Upper 0
Foliose Vulpicida pinastri JSM West Lower 0
Foliose Vulpicida pinastri JSM South Upper 0
Foliose Vulpicida pinastri JSM South Lower 0
Foliose Vulpicida pinastri JSM North Upper 0
Foliose Vulpicida pinastri JSM North Lower 0
Foliose Vulpicida pinastri JSM East Upper 0
Foliose Vulpicida pinastri JSM East Lower 0
60


Morphology Species Site Aspect Elevation Count
Foliose Vulpicida tilesii VQS West Upper 0
Foliose Vulpicida tilesii VQS West Lower 0
Foliose Vulpicida tilesii VQS South Upper 0
Foliose Vulpicida tilesii VQS South Lower 0
Foliose Vulpicida tilesii VQS North Upper 0
Foliose Vulpicida tilesii VQS North Lower 2
Foliose Vulpicida tilesii VQS East Upper 1
Foliose Vulpicida tilesii VQS East Lower 0

Foliose Vulpicida tilesii PIK West Upper 1
Foliose Vulpicida tilesii PIK West Lower 5
Foliose Vulpicida tilesii PIK South Upper 0
Foliose Vulpicida tilesii PIK South Lower 0
Foliose Vulpicida tilesii PIK North Upper 0
Foliose Vulpicida tilesii PIK North Lower 0
Foliose Vulpicida tilesii PIK East Upper 0
Foliose Vulpicida tilesii PIK East Lower 0

Foliose Vulpicida tilesii GLA West Upper 0
Foliose Vulpicida tilesii GLA West Lower 0
Foliose Vulpicida tilesii GLA South Upper 0
Foliose Vulpicida tilesii GLA South Lower 0
Foliose Vulpicida tilesii GLA North Upper 0
Foliose Vulpicida tilesii GLA North Lower 0
Foliose Vulpicida tilesii GLA East Upper N/A
Foliose Vulpicida tilesii GLA East Lower N/A

Foliose Vulpicida tilesii JSM West Upper 0
Foliose Vulpicida tilesii JSM West Lower 0
Foliose Vulpicida tilesii JSM South Upper 0
Foliose Vulpicida tilesii JSM South Lower 0
Foliose Vulpicida tilesii JSM North Upper 0
Foliose Vulpicida tilesii JSM North Lower 0
Foliose Vulpicida tilesii JSM East Upper 0
Foliose Vulpicida tilesii JSM East Lower 0
61


Morphology Species Site Aspect Elevation Count
Foliose Xanthoparmelia chlorochroa VQS West Upper 25
Foliose Xanthoparmelia chlorochroa VQS West Lower 1
Foliose Xanthoparmelia chlorochroa VQS South Upper 0
Foliose Xanthoparmelia chlorochroa VQS South Lower 4
Foliose Xanthoparmelia chlorochroa VQS North Upper 1
Foliose Xanthoparmelia chlorochroa VQS North Lower 9
Foliose Xanthoparmelia chlorochroa VQS East Upper 1
Foliose Xanthoparmelia chlorochroa VQS East Lower 0

Foliose Xanthoparmelia chlorochroa PIK West Upper 5
Foliose Xanthoparmelia chlorochroa PIK West Lower 0
Foliose Xanthoparmelia chlorochroa PIK South Upper 0
Foliose Xanthoparmelia chlorochroa PIK South Lower 9
Foliose Xanthoparmelia chlorochroa PIK North Upper 2
Foliose Xanthoparmelia chlorochroa PIK North Lower 6
Foliose Xanthoparmelia chlorochroa PIK East Upper 1
Foliose Xanthoparmelia chlorochroa PIK East Lower 1

Foliose Xanthoparmelia chlorochroa GLA West Upper 3
Foliose Xanthoparmelia chlorochroa GLA West Lower 14
Foliose Xanthoparmelia chlorochroa GLA South Upper 23
Foliose Xanthoparmelia chlorochroa GLA South Lower 17
Foliose Xanthoparmelia chlorochroa GLA North Upper 31
Foliose Xanthoparmelia chlorochroa GLA North Lower 2
Foliose Xanthoparmelia chlorochroa GLA East Upper N/A
Foliose Xanthoparmelia chlorochroa GLA East Lower N/A

Foliose Xanthoparmelia chlorochroa JSM West Upper 25
Foliose Xanthoparmelia chlorochroa JSM West Lower 29
Foliose Xanthoparmelia chlorochroa JSM South Upper 39
Foliose Xanthoparmelia chlorochroa JSM South Lower 0
Foliose Xanthoparmelia chlorochroa JSM North Upper 55
Foliose Xanthoparmelia chlorochroa JSM North Lower 21
Foliose Xanthoparmelia chlorochroa JSM East Upper 0
Foliose Xanthoparmelia chlorochroa JSM East Lower 0
62


Morphology Species Site Aspect Elevation Count
Foliose Xanthoparmelia cumberlandia VQS West Upper 35
Foliose Xanthoparmelia cumberlandia VQS West Lower 55
Foliose Xanthoparmelia cumberlandia VQS South Upper 41
Foliose Xanthoparmelia cumberlandia VQS South Lower 20
Foliose Xanthoparmelia cumberlandia VQS North Upper 11
Foliose Xanthoparmelia cumberlandia VQS North Lower 17
Foliose Xanthoparmelia cumberlandia VQS East Upper 14
Foliose Xanthoparmelia cumberlandia VQS East Lower 2

Foliose Xanthoparmelia cumberlandia PIK West Upper 0
Foliose Xanthoparmelia cumberlandia PIK West Lower 0
Foliose Xanthoparmelia cumberlandia PIK South Upper 1
Foliose Xanthoparmelia cumberlandia PIK South Lower 0
Foliose Xanthoparmelia cumberlandia PIK North Upper 5
Foliose Xanthoparmelia cumberlandia PIK North Lower 6
Foliose Xanthoparmelia cumberlandia PIK East Upper 97
Foliose Xanthoparmelia cumberlandia PIK East Lower 94

Foliose Xanthoparmelia cumberlandia GLA West Upper 31
Foliose Xanthoparmelia cumberlandia GLA West Lower 31
Foliose Xanthoparmelia cumberlandia GLA South Upper 4
Foliose Xanthoparmelia cumberlandia GLA South Lower 23
Foliose Xanthoparmelia cumberlandia GLA North Upper 4
Foliose Xanthoparmelia cumberlandia GLA North Lower 3
Foliose Xanthoparmelia cumberlandia GLA East Upper N/A
Foliose Xanthoparmelia cumberlandia GLA East Lower N/A

Foliose Xanthoparmelia cumberlandia JSM West Upper 17
Foliose Xanthoparmelia cumberlandia JSM West Lower 14
Foliose Xanthoparmelia cumberlandia JSM South Upper 18
Foliose Xanthoparmelia cumberlandia JSM South Lower 268
Foliose Xanthoparmelia cumberlandia JSM North Upper 253
Foliose Xanthoparmelia cumberlandia JSM North Lower 96
Foliose Xanthoparmelia cumberlandia JSM East Upper 43
Foliose Xanthoparmelia cumberlandia JSM East Lower 2
63


Morphology Species Site Aspect Elevation Count
Foliose Xanthoparmelia wyomingica VQS West Upper 67
Foliose Xanthoparmelia wyomingica VQS West Lower 20
Foliose Xanthoparmelia wyomingica VQS South Upper 38
Foliose Xanthoparmelia wyomingica VQS South Lower 63
Foliose Xanthoparmelia wyomingica VQS North Upper 28
Foliose Xanthoparmelia wyomingica VQS North Lower 32
Foliose Xanthoparmelia wyomingica VQS East Upper 5
Foliose Xanthoparmelia wyomingica VQS East Lower 3

Foliose Xanthoparmelia wyomingica PIK West Upper 13
Foliose Xanthoparmelia wyomingica PIK West Lower 113
Foliose Xanthoparmelia wyomingica PIK South Upper 25
Foliose Xanthoparmelia wyomingica PIK South Lower 37
Foliose Xanthoparmelia wyomingica PIK North Upper 12
Foliose Xanthoparmelia wyomingica PIK North Lower 10
Foliose Xanthoparmelia wyomingica PIK East Upper 12
Foliose Xanthoparmelia wyomingica PIK East Lower 0

Foliose Xanthoparmelia wyomingica GLA West Upper 6
Foliose Xanthoparmelia wyomingica GLA West Lower 53
Foliose Xanthoparmelia wyomingica GLA South Upper 13
Foliose Xanthoparmelia wyomingica GLA South Lower 31
Foliose Xanthoparmelia wyomingica GLA North Upper 16
Foliose Xanthoparmelia wyomingica GLA North Lower 2
Foliose Xanthoparmelia wyomingica GLA East Upper N/A
Foliose Xanthoparmelia wyomingica GLA East Lower N/A

Foliose Xanthoparmelia wyomingica JSM West Upper 67
Foliose Xanthoparmelia wyomingica JSM West Lower 80
Foliose Xanthoparmelia wyomingica JSM South Upper 13
Foliose Xanthoparmelia wyomingica JSM South Lower 0
Foliose Xanthoparmelia wyomingica JSM North Upper 14
Foliose Xanthoparmelia wyomingica JSM North Lower 10
Foliose Xanthoparmelia wyomingica JSM East Upper 0
Foliose Xanthoparmelia wyomingica JSM East Lower 0
64


Morphology Species Site Aspect Elevation Count
Foliose Xanthoria elegans VQS West Upper 179
Foliose Xanthoria elegans VQS West Lower 275
Foliose Xanthoria elegans VQS South Upper 236
Foliose Xanthoria elegans VQS South Lower 154
Foliose Xanthoria elegans VQS North Upper 106
Foliose Xanthoria elegans VQS North Lower 27
Foliose Xanthoria elegans VQS East Upper 161
Foliose Xanthoria elegans VQS East Lower 16

Foliose Xanthoria elegans PIK West Upper 133
Foliose Xanthoria elegans PIK West Lower 11
Foliose Xanthoria elegans PIK South Upper 321
Foliose Xanthoria elegans PIK South Lower 147
Foliose Xanthoria elegans PIK North Upper 238
Foliose Xanthoria elegans PIK North Lower 180
Foliose Xanthoria elegans PIK East Upper 79
Foliose Xanthoria elegans PIK East Lower 27

Foliose Xanthoria elegans GLA West Upper 57
Foliose Xanthoria elegans GLA West Lower 121
Foliose Xanthoria elegans GLA South Upper 118
Foliose Xanthoria elegans GLA South Lower 132
Foliose Xanthoria elegans GLA North Upper 63
Foliose Xanthoria elegans GLA North Lower 145
Foliose Xanthoria elegans GLA East Upper N/A
Foliose Xanthoria elegans GLA East Lower N/A

Foliose Xanthoria elegans JSM West Upper 208
Foliose Xanthoria elegans JSM West Lower 181
Foliose Xanthoria elegans JSM South Upper 194
Foliose Xanthoria elegans JSM South Lower 444
Foliose Xanthoria elegans JSM North Upper 163
Foliose Xanthoria elegans JSM North Lower 166
Foliose Xanthoria elegans JSM East Upper 100
Foliose Xanthoria elegans JSM East Lower 1
65


C. SITE TABLES
66


VOS
West Upper South Upper North Upper East Upper Totals
Species Total Species Total Species Total Species Total
A. contigua 76 A. contigua 57 A. contigua 60 A. contigua 65 258
A. caesiocinerea 131 A. caesiocinerea 208 A. caesiocinerea 88 A. caesiocinerea 146 573
D. musconim 0 D. musconim 3 D. musconim 0 D. musconim 0 3
L. rupicola 23 L. rupicola 42 L. rupicola 44 L. rupicola 29 138
L. atrobrunnea 119 L. atrobrunnea 177 L. atrobrunnea 191 L. atrobrunnea 140 627
L. tessellata 20 L. tessellata 42 L. tessellata 34 L. tessellata 38 134
P. minuscula 4 P. minuscula 0 P. minuscula 5 P. minuscula 0 9
R. geographicum 63 R. geographicum 28 R. geographicum 63 R. geographicum 30 184
R. chrvsoleuca 4 R. chrvsoleuca 3 R. chysoleuca 41 R. chysoleuca 23 71
T. siibiiliformis 0 T. siibiiliformis 1 T. siibiiliformis 0 T. siibiiliformis 0 1
U. americana 0 U. americana 0 U. americana 0 U. americana 0 0
U. decussata 197 U. decussata 169 U. decussata 108 U. decussata 3 477
U. virginis 473 U. virginis 442 U. virginis 265 U. virginis 200 1380
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri 0 0
V. tilesii 0 V. tilesii 0 V. tilesii 0 V. tilesii 1 1
X. chlorochroa 25 X. chlorochroa 0 X. chlorochroa 1 X. chlorochroa 1 27
X. cumberlandia 35 X. cumberlandia 41 X. cumberlandia 11 X. cumberlandia 14 101
X. wyomingica 67 X. wyomingica 38 X. wyomingica 28 X. wyomingica 5 138
X. elegans 179 X. elegans 236 X. elegans 106 X. elegans 161 682
Total 1416 Total 1487 Total 1045 Total 856 4804
West Lower South Lower North Lower East Lower Totals
Species Total Species Total Species Total Species Total
A. contigua 95 A. contigua 120 A. contigua 91 A. contigua 57 363
A. caesiocinerea 75 A. caesiocinerea 140 A. caesiocinerea 57 A. caesiocinerea 88 360
D. musconim 0 D. musconim 17 D. musconim 0 D. musconim 0 17
L. rupicola 32 L. rupicola 26 L. rupicola 56 L. rupicola 30 144
L. atrobrunnea 87 L. atrobrunnea 190 L. atrobrunnea 104 L. atrobrunnea 98 479
L. tessellata 35 L. tessellata 44 L. tessellata 17 L. tessellata 7 103
P. minuscula 4 P. minuscula 6 P. minuscula 1 P. minuscula 0 11
R. geographicum 3 R. geographicum 27 R. geographicum 27 R. geographicum 7 64
R. chysoleuca 4 R. chysoleuca 31 R. chysoleuca 9 R. chysoleuca 6 50
T. siibiiliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 0
U. americana 3 U. americana 0 U. americana 0 U. americana 0 3
U. decussata 63 U. decussata 202 U. decussata 21 U. decussata 8 294
U. virginis 304 U. virginis 333 U. virginis 75 U. virginis 40 752
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri 0 0
V. tilesii 0 V. tilesii 0 V. tilesii 2 V. tilesii 0 2
X. chlorochroa 1 X. chlorochroa 4 X. chlorochroa 9 X. chlorochroa 0 14
X. cumberlandia 55 X. cumberlandia 20 X. cumberlandia 17 X. cumberlandia 2 94
X. wyomingica 20 X. wyomingica 63 X. wyomingica 32 X. wyomingica 3 118
X. elegans 275 X. elegans 154 X. elegans 27 X. elegans 16 472
Totals 1056 1377 545 362 3340
67


PIK
West Upper South Upper North Upper East Upper Totals
Species Total Species Total Species Total Species Total
A. contigua 97 A. contigua 127 A. contigua 85 A. contigua 278 587
A. caesiocinerea 15 A. caesiocinerea 220 A. caesiocinerea 168 A. caesiocinerea 227 630
D. musconim 0 D. muscorum 0 D. musconim 0 D. muscorum 0 0
L. rupicola 157 L. rupicola 125 L. rupicola 93 L. rupicola 232 607
L. atrobnmnea 151 L. atrobnmnea 314 L. atrobnmnea 150 L. atrobnmnea 356 971
L. tessellata 29 L. tessellata 94 L. tessellata 93 L. tessellata 133 349
P. minuscula 23 P. minuscula 78 P. minuscula 6 P. minuscula 82 189
R. geographicum 351 R. geographicum 450 R. geographicum 124 R. geographicum 648 1573
R. chrysoleuca 87 R. chrysoleuca 63 R. chiysoleuca 50 R. chiysoleuca 63 263
T. subuliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 0
U. americana 11 U. americana 3 U. americana 0 U. americana 5 19
U. decussata 115 U. decussata 185 U. decussata 48 U. decussata 132 480
U. virginis 309 U. virginis 514 U. virginis 326 U. virginis 512 1661
V. pinastri 0 V. pinastri 3 V. pinastri 0 V. pinastri 0 3
V. tilesii 1 V. tilesii 0 V. tilesii 0 V. tilesii 0 1
X. chlorochroa 5 X. chlorochroa 0 X. chlorochroa 2 X. chlorochroa 1 8
X. camberlandia 0 X. cumberlandia 1 X. cumberlandia 5 X. cumberlandia 97 103
X. wyomingica 13 X. wyomingica 25 X. wyomingica 12 X. wyomingica 12 62
X. elegans 133 X. elegans 321 X. elegans 238 X. elegans 79 771
Total 1497 Total 2523 Total 1400 Total 2857 8277
West Lower South Lower North Lower East Lower Totals
Species Total Species Total Species Total Species Total
A. contigua 101 A. contigua 52 A. contigua 141 A. contigua 275 569
A. caesiocinerea 9 A. caesiocinerea 29 A. caesiocinerea 182 A. caesiocinerea 46 266
D. musconim 22 D. muscorum 6 D. musconim 0 D. muscorum 1 29
L. rupicola 91 L. rupicola 61 L. rupicola 84 L. rupicola 113 349
L. atrobnmnea 191 L. atrobnmnea 124 L. atrobnmnea 151 L. atrobnmnea 178 644
L. tessellata 37 L. tessellata 53 L. tessellata 110 L. tessellata 87 287
P. minuscula 22 P. minuscula 11 P. minuscula 24 P. minuscula 0 57
R. geographicum 222 R. geographicum 48 R. geographicum 201 R. geographicum 316 787
R. chiysoleuca 46 R. chiysoleuca 52 R. chiysoleuca 54 R. chiysoleuca 22 174
T. subuliformis 3 T. subuliformis 2 T. subuliformis 0 T. subuliformis 0 5
U. americana 1 U. americana 0 U. americana 0 U. americana 0 1
U. decussata 82 U. decussata 24 U. decussata 100 U. decussata 1 207
U. virginis 256 U. virginis 337 U. virginis 240 U. virginis 28 861
V. pinastri 1 V. pinastri 0 V. pinastri 0 V. pinastri 0 1
V. tilesii 5 V. tilesii 0 V. tilesii 0 V. tilesii 0 5
X. chlorochroa 0 X. chlorochroa 9 X. chlorochroa 6 X. chlorochroa 1 16
X. cumberlandia 0 X. cumberlandia 0 X. cumberlandia 6 X. cumberlandia 94 100
X. wyomingica 113 X. wyomingica 37 X. wyomingica 10 X. wyomingica 0 160
X. elegans 11 X. elegans 147 X. elegans 180 X. elegans 27 365
Total 1213 Total 992 Total 1489 Total 1189 4883
68


GLA
West Upper South Upper North Upper East Upper Totals
Species Total Species Total Species Total Species Total
A. contigua 131 A. contigua 201 A. contigua 157 A. contigua N/A 489
A. caesiocinerea 42 A. caesiocinerea 140 A. caesiocinerea 44 A. caesiocinerea N/A 226
D. muscorum 1 D. muscorum 3 D. muscorum 1 D. muscorum N/A 5
L. rupicola 140 L. rupicola 246 L. rupicola 118 L. rupicola N/A 504
L. atrobrunnea 489 L. atrobrunnea 627 L. atrobrunnea 349 L. atrobrunnea N/A 1465
L. tessellata 102 L. tessellata 185 L. tessellata 126 L. tessellata N/A 413
P. minuscula 40 P. minuscula 50 P. minuscula 5 P. minuscula N/A 95
R. geographicum 718 R. geographicum 869 R. geographicum 307 R. geographicum N/A 1894
R. chrysoleuca 55 R. chrysoleuca 133 R. chrysoleuca 50 R. chrysoleuca N/A 238
T. subuliformis 0 T. subuliformis 1 T. subuliformis 0 T. subuliformis N/A 1
U. americana 12 U. americana 71 U. americana 32 U. americana N/A 115
U. decussata 347 U. decussata 410 U. decussata 371 U. decussata N/A 1128
U. Virginia 643 U. virginis 1037 U. virginis 225 U. virginis N/A 1905
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri N/A 0
V. tilesii 0 V. tilesii 0 V. tilesii 0 V. tilesii N/A 0
X. chlorochroa 3 X. chlorochroa 23 X. chlorochroa 31 X. chlorochroa N/A 57
X. cumberlandia 31 X. cumberlandia 4 X. cumberlandia 4 X. cumberlandia N/A 39
X. wyomingica 6 X. wyomingica 13 X. wyomingica 16 X. wyomingica N/A 35
X. elegans 57 X. elegans 118 X. elegans 63 X. elegans N/A 238
Total 2817 Total 4131 Total 1899 Total 8847
West Uower South Uower North Uower East Uower Totals
Species Total Species Total Species Total Species Total
A. contigua 198 A. contigua 236 A. contigua 210 A. contigua N/A 644
A. caesiocinerea 122 A. caesiocinerea 137 A. caesiocinerea 258 A. caesiocinerea N/A 517
D. muscorum 17 D. muscorum 1 D. muscorum 4 D. muscorum N/A 22
L. rupicola 185 L. rupicola 144 L. rupicola 133 L. rupicola N/A 462
L. atrobrunnea 505 L. atrobrunnea 517 L. atrobrunnea 411 L. atrobrunnea N/A 1433
L. tessellata 187 L. tessellata 162 L. tessellata 223 L. tessellata N/A 572
P. minuscula 18 P. minuscula 37 P. minuscula 12 P. minuscula N/A 67
R. geographicum 579 R. geographicum 633 R. geographicum 398 R. geographicum N/A 1610
R. chrysoleuca 114 R. chrysoleuca 105 R. chrysoleuca 11 R. chrysoleuca N/A 230
T. subuliformis 0 T. subuliformis 0 T. subuliformis 0 T. subuliformis N/A 0
U. americana 7 U. americana 43 U. americana 52 U. americana N/A 102
U. decussata 327 U. decussata 76 U. decussata 150 U. decussata N/A 553
U. virginis 865 U. virginis 436 U. virginis 369 U. virginis N/A 1670
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri N/A 0
V. tilesii 0 V. tilesii 0 V. tilesii 0 V. tilesii N/A 0
X. chlorochroa 14 X. chlorochroa 17 X. chlorochroa 2 X. chlorochroa N/A 33
X. cumberlandia 31 X. cumberlandia 23 X. cumberlandia 3 X. cumberlandia N/A 57
X. wyomingica 53 X. wyomingica 31 X. wyomingica 2 X. wyomingica N/A 86
X. elegans 121 X. elegans 132 X. elegans 145 X. elegans N/A 398
Total 3343 Total 2730 Total 2383 Total 8456
69


JSM
West Upper South Upper North Upper East Upper Totals
Species Total Species Total Species Total Species Total
A. contigua 258 A. contigua 407 A. contigua 496 A. contigua 317 1478
A. caesiocinerea 226 A. caesiocinerea 350 A. caesiocinerea 155 A. caesiocinerea 126 857
D. musconim 18 D. muscorum 5 D. musconim 9 D. muscorum 0 32
L. rupicola 70 L. rupicola 71 L. rupicola 127 L. rupicola 47 315
L. atrobninnea 235 L. atrobninnea 256 L. atrobninnea 285 L. atrobninnea 189 965
L. tessellata 0 L. tessellata 0 L. tessellata 50 L. tessellata 35 85
P. minuscula 35 P. minuscula 7 P. minuscula 0 P. minuscula 42 84
R. geographicum 125 R. geographicum 185 R. geographicum 174 R. geographicum 232 716
R. chrvsoleuca 47 R. chrvsoleuca 10 R. chiysoleuca 49 R. chiysoleuca 0 106
T. subuliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 T. siibiiliformis 0 0
U. americana 47 U. americana 8 U. americana 9 U. americana 0 64
U. decussata 24 U. decussata 0 U. decussata 0 U. decussata 0 24
U. virginis 283 U. virginis 145 U. virginis 57 U. virginis 0 485
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri 0 0
V. tilesii 0 V. tilesii 0 V. tilesii 0 V. tilesii 0 0
X. chlorochroa 25 X. chlorochroa 39 X. chlorochroa 55 X. chlorochroa 0 119
X. cumberlandia 17 X. cumberlandia 18 X. cumberlandia 253 X. cumberlandia 43 331
X. wyomingica 67 X. wyomingica 13 X. wyomingica 14 X. wyomingica 0 94
X. elegans 208 X. elegans 194 X. elegans 163 X. elegans 100 665
Total 1685 Total 1708 Total 1896 Total 1131 6420
West Uower South Uower North Uower East Uower Totals
Species Total Species Total Species Total Species Total
A. contigua 323 A. contigua 439 A. contigua 465 A. contigua 29 1256
A. caesiocinerea 205 A. caesiocinerea 103 A. caesiocinerea 201 A. caesiocinerea 2 511
D. musconim 5 D. muscorum 0 D. musconim 0 D. muscorum 0 5
L. rupicola 62 L. rupicola 10 L. rupicola 139 L. rupicola 5 216
L. atrobninnea 179 L. atrobrunnea 279 L. atrobrunnea 352 L. atrobrunnea 1 811
L. tessellata 0 L. tessellata 0 L. tessellata 0 L. tessellata 6 6
P. minuscula 3 P. minuscula 0 P. minuscula 5 P. minuscula 0 8
R. geographicum 118 R. geographicum 5 R. geographicum 426 R. geographicum 31 580
R. chiysoleuca 92 R. chiysoleuca 82 R. chiysoleuca 69 R. chiysoleuca 0 243
T. subuliformis 0 T. subuliformis 0 T. subuliformis 0 T. subuliformis 0 0
U. americana 34 U. americana 4 U. americana 24 U. americana 0 62
U. decussata 5 U. decussata 5 U. decussata 17 U. decussata 0 27
U. virginis 184 U. virginis 81 U. virginis 78 U. virginis 0 343
V. pinastri 0 V. pinastri 0 V. pinastri 0 V. pinastri 0 0
V. tilesii 0 V. tilesii 0 V. tilesii 0 V. tilesii 0 0
X. chlorochroa 29 X. chlorochroa 0 X. chlorochroa 21 X. chlorochroa 0 50
X. cumberlandia 14 X. cumberlandia 268 X. cumberlandia 96 X. cumberlandia 2 380
X. wyomingica 80 X. wyomingica 0 X. wyomingica 10 X. wyomingica 0 90
X. elegans 181 X. elegans 444 X. elegans 166 X. elegans 1 792
Total 1514 Total 1720 Total 2069 Total 77 5380
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D. SITE TOTALS
VQS Totals PIK Totals
Acarospora contigua 621 Acarospora contigua 1156
Aspicilia caesiocinerea 933 Aspicilia caesiocinerea 896
Diploschistes muscorum 20 Diploschistes muscorum 29
Lecanora rupicola 282 Lecanora rupicola 956
Lecidea atrobrunnea 1106 Lecidea atrobrunnea 1615
Lecidea tessellata 237 Lecidea tessellata 636
Pseudephebe minuscula 20 Pseudephebe minuscula 246
Rhizocarpon geographicum 248 Rhizocarpon geographicum 2360
Rhizoplaca chrysoleuca 121 Rhizoplaca chrysoleuca 437
Thamnolia subuliformis 1 Thamnolia subuliformis 5
Umbilicaria americana 3 Umbilicaria americana 20
Umbilicaria decussata 111 Umbilicaria decussata 687
Umbilicaria virginis 2132 Umbilicaria virginis 2522
Vulpicida pinastri 0 Vulpicida pinastri 4
Vulpicida tilesii 3 Vulpicida tilesii 6
Xanthoparmelia chlorochroa 41 Xanthoparmelia chlorochroa 24
Xanthoparmelia cumberlandia 195 Xanthoparmelia cumberlandia 203
Xanthoparmelia wyomingica 256 Xanthoparmelia wyomingica 222
Xanthoria elegans 1154 Xanthoria elegans 1136
Site Total 8144 Site Total 13160
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GLA Totals JSM Totals
Acarospora contigua 1133 Acarospora contigua 2734
Aspicilia caesiocinerea 743 Aspicilia caesiocinerea 1368
Diploschistes muscorum 27 Diploschistes muscorum 37
Lecanora rupicola 966 Lecanora rupicola 531
Lecidea atrobrunnea 2898 Lecidea atrobrunnea 1776
Lecidea tessellata 985 Lecidea tessellata 91
Pseudephebe minuscula 162 Pseudephebe minuscula 92
Rhizocarpon geographicum 3504 Rhizocarpon geographicum 1296
Rhizoplaca chrysoleuca 468 Rhizoplaca chrysoleuca 349
Thamnolia subuliformis 1 Thamnolia subuliformis 0
Umbilicaria americana 217 Umbilicaria americana 126
Umbilicaria decussata 1681 Umbilicaria decussata 51
Umbilicaria virginis 3575 Umbilicaria virginis 828
Vulpicida pinastri 0 Vulpicida pinastri 0
Vulpicida tilesii 0 Vulpicida tilesii 0
Xanthoparmelia chlorochroa 90 Xanthoparmelia chlorochroa 169
Xanthoparmelia cumberlandia 96 Xanthoparmelia cumberlandia 711
Xanthoparmelia wyomingica 121 Xanthoparmelia wyomingica 184
Xanthoria elegans 636 Xanthoria elegans 1457
Site Total 17303 Site Total 11800
72


E. SPECIES PHOTOGRAPHS
73


Acarosyora contisua
74


Asyicilia caesiocinerea
75


Diyloschistes muscorum
76


Lecanora ruyicola
11


Lecidea atrobrunnea
78


Lecidea tessellate
79


Pseudeyhebe minuscula
80


Rhizocarpon seosrayhicum
81


Rhizovlaca chrysoleuca
82


Thamnolia subuliformis
83


Umbilicaria Americana
84


Umbilicaria decussate
85


Umbilicaria virsinis
86


Vulyicida yinastri
87


Vulyicida tilesii
88


Xanthoyarmelia chlorochroa
89


Xanthoyarmelia cumberlandia
90


Xanthoyarmelia wyominsica
91


Xanthoria elesans
92


F. GOOGLE EARTH HISTORICAL IMAGERY
Google Earth imagery of the study sites October 2013.
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Full Text

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A GEOGRAPHICAL ANALYSIS OF ALPINE LICHEN IN ROCKY MOUNTAIN NATIONAL PARK by ERIK DOUGLAS AHL B.S., Western Carolina University, 2011 A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of t he requirements for the degree of Master of Science Environmental Science s program 2016

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ii 2016 ERIK DOUGLAS AHL ALL RIGHTS RESERVED

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iii This thesis for the Master of Science degree by Erik Douglas Ahl has been approved for the Enviro nmental Sciences Program by Casey Allen, Chair Frederick Chambers David Knochel December 17 th 2016

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iv Ahl, Erik Douglas (MS, Environmental Sciences) A Geographical Analysis of Alpine Lichen in Rocky Mountain National Park Thesis directed by Profes sor Casey Allen ABSTRACT In high alpine ecosystems, landscapes often seem barren to the naked eye due to intense exposure from UV light, temperature fluxes, and desiccation. Species present are only those resilient enough to have adapted to such harsh cond itions, and these are usually ground hugging species such as those in Biological Soil Crusts (BSCs). Consisting of various species of cyanobacteria, algae, lichens, and mosses, BSCs have been studied for decades in dry land environments all over the world, yet little BSC research focuses on high altitude ecosystems. This study set out to analyze BSC distribution and characteristics. In situ however, lichens dominated rocks and soil at the chosen sites and thus became the focus for the study. The four sites (three high alpine, one subalpine) utilized were already under study in Rocky Mountain National Park (RMNP) with the Rocky Mountain Inventory and Monitoring Network (ROMN) in conjunction with the Global Observation Initiative in Alpine Environments (GLORI A). Implementing field techniques outlined by GLORIA, species were photographed, identified, inventoried and then analyzed using SPSS statistical software. It was suspected that findings would show correlations between aspects, soil, and rock types. Relati onships between some species and specific rock types, such as siliceous and granitic, were initially discovered, as well as variability in types among other species.

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v The form and content of this abstract are approved. I recommend its publication Approved: Casey Allen

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vi TABLE OF CONTENTS CHAPTER I ................................ ................................ ................................ ................................ .... 1 I. INTRODUCTION ................................ ................................ ................................ ................................ 1 1.1 Internal Function ................................ ................................ ................................ ............................. 2 1.2 Classification ................................ ................................ ................................ ................................ ... 2 1.3 Effects on Environment ................................ ................................ ................................ .................. 4 1.4 Lichen Distribution ................................ ................................ ................................ ......................... 5 CHAPTER II ................................ ................................ ................................ ................................ ... 8 II. SITE SETTING ................................ ................................ ................................ ................................ .... 8 CHAPTER III ................................ ................................ ................................ ............................... 10 III. METHODS ................................ ................................ ................................ ................................ ....... 10 3.1 Field Work ................................ ................................ ................................ ................................ .... 10 3.2 Post Field Work ................................ ................................ ................................ ............................ 11 3.3 Statistical Analysis ................................ ................................ ................................ ........................ 13 CHAPTER IV ................................ ................................ ................................ ............................... 14 IV. RESULTS ................................ ................................ ................................ ................................ ......... 14 4.1 VQS ................................ ................................ ................................ ................................ ............... 14 4.2 PIK ................................ ................................ ................................ ................................ ................ 15 4.3 GLA ................................ ................................ ................................ ................................ .............. 16 4.4 JSM ................................ ................................ ................................ ................................ ............... 18 ................................ ................................ ................................ ....... 20 ................................ ................................ ................................ ...................... 21 4.7 Rank Abundance ................................ ................................ ................................ ........................... 23 4.8 SPSS ................................ ................................ ................................ ................................ .............. 26 CHAPTER V ................................ ................................ ................................ ................................ 32 V. DISCUSSION ................................ ................................ ................................ ................................ .... 32 CHAPTER VI ................................ ................................ ................................ ............................... 34 VI. CONCLUSION ................................ ................................ ................................ ................................ 34 REFERENCES ................................ ................................ ................................ ............................. 37

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vii APPENDIX ................................ ................................ ................................ ................................ ... 43 A. LICHEN LIST FROM RMNP ................................ ................................ ................................ ........... 43 B. SPECIES TABLES ................................ ................................ ................................ ............................ 47 C. SITE TABLES ................................ ................................ ................................ ................................ ... 66 D. SITE TOTALS ................................ ................................ ................................ ................................ ... 71 E. SPECIES PHOTOGRAPHS ................................ ................................ ................................ .............. 73 F. GOOGLE EARTH HISTORICAL IMAGERY ................................ ................................ ................. 93

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1 CHAPTER I INTRODUCTION Somewhere amongst macro and microorganisms, splayed acr oss soil, rocks, and trees are often brightly colored plant like creatures called lichens. From small patches to large carpet like spreads, these often overlooked non vascular organisms can be found throughout the world and across all ecosystems: from the hot deserts, to cold alpine tundra, rainforests, coastal systems and even Antarctica. Lichens are part of a larger collection of species, along with algae and mosses that can be classified under the umbrella of Biological Soil Crusts (BSCs) (Belnap and Lan ge 2003b, Belnap et al. 2001, Purvis 2000) Though much has been discovered in recent decades regarding lichens, most studies have focused on lower elevation species, centering specifically in hyper arid and arid deserts (Allen 2005, Allen 2010) Excludin g a handful of studies conducted outside of the United States (Breen and Lvesque 2008, Pohl et al. 2009, Karsten, Ltz and Holzinger 2010, Yoshitake et al. 2010) little research has been conducted in regards to higher elevation lichen in the United States With this in mind, this study aims to shed more light on under studied alpine lichen in the United States. Taking place in Rocky Mountain National Park (RMNP), Colorado, four sites were selected as proxies for determining richness, abundance, and cover of lichen species at differing elevations, and teasing out their locational relationships in regards to aspect, elevation, and site location to investigate perceived differences between alpine and subalpine (Belnap and Lange 2003a, Brodo, Sharnoff and Shar noff 2001b)

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2 First off, a brief discussion of the lichens internal function (1.1) and classification (1.2) will be offered, followed by their role in the environment (1.3) Then a description of planation of methods implemented in the field and for the post field work (III). Finally, results (IV) are presented and analyses are outlined along with an in depth discussion ( V) of those techniques and results leading to a brief conclusion (V I ). 1.1 In ternal Function Though numerous vascular and nonvascular plant species inhabit these high elevation landscapes, the predominant species found sprawling across the rocks and soils in often bright contrast are lichens. Carl Linnaeus, the father of taxonomy, is said to have called Lichen ) to encompass all lichens (Walewski 2007) Little did he realize that lichens are not so easily classified into any mplex relationship developed between two separate organisms working together for a mutually beneficial relationship. This symbiosis is the marriage between a fungi and a photosynthetic partner of algae or cyanobacteria (Brodo, Sharnoff and Sharnoff 2001c) a partnership that allows for the algae to live within the body of the fungus, while the fungus consumes algal cells for its own nourishment. These algal cells replenish at a rate much quicker than they are consumed, allowing for a very efficient form of parasitism in which both parties benefit (McCune and Goward 1995) 1.2 Classification There are approximately 14,000 lichen species throughout the world, each of which varies in shape, size, and color. Generally, lichen are classified into four growth fo rms

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3 utilizing its vegetative body known as the thallus: foliose, fruticose, crustose, and squamulose (Brodo et al. 2001c) Foliose lichens, which encompassed 10 of the 19 species in this study, often portray a leafy in shape, as the word implies. These l obed appendages host a more or less flattened thallus and are securely anchored to the substrate through the use of hyphae. Hyphae are filamentous elements of the fungal component of the lichen, consisting of cells arranged in threads of varying shapes, si zes, and branching patterns. Also instrumental for anchorage to substrate by foliose lichens are rhizines branched or unbranched bottom growing appendages that often occur in tufted or fibrous bundles (Brodo et al. 2001c) Fruiticose lichens, two of the 19 species in this study, come in a more shrubby, three dimensional form with a radial symmetry in their stalks (McCune and Goward 1995) Growing in a more erect or pendent fashion, they resemble foliose lichens, but their thali are comprised in a more th ree than two dimensional fashion (Brodo et al. 2001c) Crustose lichens resemble, as the name implies, a crust across the chosen surface substrate and were represented in this study by seven of the 19 spe cies. Found in a vast array of radiant colors, the se lichens grow on and among the particles of the substrate forming small to very large patches that can be quite thick or rough (Brodo et al. 2001c) What differentiates crustose from the foliose and fruticose is the absence of a lower cortex, or lower fu ngal layer. Squamulose lichen, which were not represented in the study sites, consist of a cross between foliose and crustose in both growth and formation (Brodo et al. 2001c) These

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4 growth forms are composed of clusters in small roundish flakes and come in a various shapes with slightly lobate margins (McCune and Goward 1995) 1.3 Effects on Environment Lichens play significant roles with their surrounding biota (Kar et al. 2014) For example, like most vegetative species, lichens no matter their locat ion, from dese rt plains to alpine ecosystems help with C sequestration and N fixation (Belnap and Lange 2003b, Johnston 1997, Barger et al. 2006, Yoshitake et al. 2010) These types of desiccating resilient organisms are known as poikilohydric (Viles 2008) because they are able to dry out and suspend respiration without any negative side effects, while being environmentally dependent on their water (Green and Lange 1995) Exercising such environmental resilience to adverse conditions, such as those experie nced by all vegetation in the high elevations of the alpine tundra in Colorado, lichen are known to live and thrive for hundreds of years (Lalley and Viles 2007) They have the ability to colonize just about anywhere including bare rocks and soil. Lending to not only fixation and influencing hydrological patterns, their influence on the erosion of soil and rock can be paramount (Zelikova et al. 2012) The hyphae appendages extending below the thallus, and being able to bore into solid rock substrate, can al low for the speeding up of local rock decay processes (McIlroy de la Rosa, Warke and Smith 2012) This mechanical decay can also be exacerbated by chemical decay from the carbonic acid formed in lichen metabolic activities (Raggio, Green and Sancho 2015) eating away at rocks rich in calcium carbonate that they reside on (Brodo et al. 2001c)

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5 At lower elevations, lichen can also be found colonizing the exterior of trees along their bark (Ames et al. 2012) Faster growing foliose and fruticose species tend t o colonize first, with crustose species being the last to show up on tree bark. As forest inhabitants, they affect the dispersion of rainwater nutrients as they flow down the body of the tree (Brodo et al. 2001c) This same process occurs with nutrients ab sorbed from fog, thus affecting what concentrations are reflected in the ground below, affecting the surrounding plant growth. Lichen also play an important role in environmental monitoring (Zedda et al. 2011, Shukla 2014, Bosch Roig et al. 2013) as they are well known for their sensitivity to air pollution (Corbridge and Weber 1998, Kularatne and de Freitas 2013) A sensitivity, that is directly related to their uncanny ability to absorb chemicals rapidly from the air (Brodo et al. 2001c, Belnap et al. 20 01, Walewski 2007, Kularatne and de Freitas 2013) Sulfur dioxide and components of acid rain (Purvis 2000, Cuculovic et al. 2014) (sulfuric and nitric acids) are especially detrimental to lichen species. In areas of high industrial pollution, lichen tend to be absent and this can be used as a biomarker to an areas ambient pollution (Brodo, Sharnoff and Sharnoff 2001a) 1.4 Lichen Distribution Almost as varied as their biology and structure, lichens also remain a widespread organism around the world (Feuer er and Hawksworth 2007) (Brodo et al. 2001c) notes in his book that all lichen species have climatic tolerances and physiological constraints (Peksa and Skaloud 2011, Gauslaa 2014) So, with that in mind, the authors classified distribution patterns in re lation to major climatic regions across the North American continent. Their classification is broken down into eight climatic elements: Arctic Alpine, Boreal, Temperate,

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6 Central Grasslands, Western Sector, Western Montane, Madrean, Tropical, Oceanic and Ma ritime. The Arctic Alpine in the more northern parts of the continent is an arid ecosystem with harsh elements (Bonan 2010, King et al. 2012) Typically situated at elevations above 3,505 M, only the resilient species like lichen may thrive under the hars her elements (Bjerke 2008) experienced there including freezing temperatures, snow, high winds greater UV exposure and desiccation (Belnap 2006) Most lichen species residing in these elements are able to shut down internal processes during long dry spells enduring the severe frost in winter, only to spring into action once the first rain or mist lays moisture on them (Dahl 1954, Stephenson 2010, Belnap 2006) Alpine environments, as in ( Fig. 1.4.1), comprise about 20% of land cover in some areas. Within these elevations, alpine ecosystems are deemed an important source of information due to their particular vulnerability to climate change and lack of environmental studies on their ecophysiological performance (Karsten and Holzinger 2012, Belnap and Lange 2003a, Breen and Lvesque 2008) With this in mind, this lichen study was focused on these important ecosystems found within Rocky Mountain National Park, and how species abundance and diversity might contrast to a subalpine site.

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7 Figure 1.4.1 Alpine Tu ndra locations around the world (dark blue), typically above 3,300 meters. credit https://infogr.am/alpine 9177284109

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8 CHAPTER II SITE SETTING Under the scope of few studies in relation to alpine environments, as well as, the lack of lichen ba sed studies in alpine systems, the author has engaged in a study pertaining to lichens while building a biological inventory to act as baseline data for further studies, while adding to previous in situ inventories. All work is in accordance with four previously studied sites under the parks Rocky Mountain Monitoring Network (ROMN) and Global Observation Research Initiative i n Alpine Environments (GLORIA). Fig ure 2.1 Satellite image of R.M.N.P. displaying the location of the four study sites

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9 ROMN is comprised of 32 vital monitoring sites spread throughout the National Park Service under program manager Mike Britton. In conjunction with GLORIA, an internationa l monitoring network established in 2001, ROMN established these sites on RMNP in 2009 (Ashton 2011) GLORIA summit criteria, adopted for use by ROMN, outlined in their field manual (Pauli et al. 2004) was the basis for site choice. Located in the higher e levations of RMNP, GLORIA had three unnamed alpine sites and one subalpine already under monitoring (Table 2.1): (VQS, PIK, GLA) and Jack Straw Mountain (JSM), respectively. These sites are located in the southwestern part of the park by Milner Pass and ac cessed via the Ute to the Mt. Ida trail. The three alpine sites run along the continental divide in a north (VQS) to south direction, with GLA acting as a southern terminus. The subalpine site (JSM) sits just west of the ridge to Mt. Ida and was accessed b y parking at the Lake Irene trailhead, crossing trail ridge road, and bush whacking to the summit. The alpine and subalpine are typically ecosystems that are found above tree line, though this elevation changes with increased latitude. In Colorado, the el evation range is approximately 3,048 m to 3505.2 m for subalpine and 3505.2 m and higher for an alpine designation. GLORIA also calls for the sites to be similar in disturbance, in this case meaning limited park traffic and geology. All four sites have sim ilar geology consisting of granite, gneiss, and schist, with a couple instances of quartz outcrops (United States Department of Agriculture 2007) Table 2. 1 Name, location, elevation, and vegetation zones of the four sites (Asht on, 2011)

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10 CHAPTER III M ETHODS 3.1 Field Work So as not to reinvent the wheel and stay in accordance to practices implemented by GLORIA and ROMN, monitoring protocols developed by GLORIA were utilized for this study. As Ashton (2001) asserts: als of the GLORIA program are to provide a global baseline for vegetation monitoring in alpine environments to assess the risks of biodiversity loss and ecosystem instability from climate change. The methodology is extended by cooperators, such as the ROMN to create a long term monitoring network at contains components for soil condition, tree line movement, and human disturbance. The GLORIA design cal ls for four sentinel sites, sites that are all similar in geology, climate, disturbance, and land use history, such as the four used in this study. Eight summit area sections are used to measure the dimensions of the peak (area and slope) from the highest point to 10m below (Ashton 2011) GLORIA uses this method to measure ground cover that has been divided into seven classes. However, it is in the interest of this study to do a biological inventory, so quadrat clusters will be placed within the plots in 3m 2 sections, equating to 8 quadrats per peak placed in respect to aspect. This is a GLORIA standard to measure vegetation change (cf., Ashton, 2011) and will be implemented by this study for inventorying and noting disturbance effects to recorded species of lichen.

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11 Summit coordinates were supplied by GLORIA and entered into a Garmin GPS 60CSx handheld GPS unit to ensure correct site location. Each summit has been marked with a pile of rocks that also served as protection for their data collection units. 3m 2 quadrats were measured using a Komelon 6611 100ft open reel fiberglass tape measure and staked with 6.35cm x 8.89cm x 53.34cm orange wire survey flags. These flags were temporarily placed during the duration of the study and removed upon completion of tha Flags were placed into the measured corners of each quadrat, resembling a tic tac toe formation. A GPS coordinate was logged with the Garmin unit at all four outer corners of each quadrat. Each of the four aspects per site was measured ou t into upper and lower sections. The upper section was from peak to five meters below and the lower section was between the five meter mark and 10 meters from peak. A 3m 2 quadrat was measured out in random placement within the confines of the upper and low er measurements of each aspect. Photographs were taken using a Nikon D80 10.2 MP camera for documentation of in situ species identification. All photographic data along with site appropriate notes were recorded in a Rite in the Rain field j ournal and site information was written and photographed on a dry erase board. 3.2 Post Field Work The PI and advisor reviewed the photographs and using a dichotomous key in the book Lichens of North America (Brodo et al. 2001c) identified each of the species represent ed. Once identified, each image was then checked against a list of species that RMNP has published on a website (Appendix A ),as well as double checked in a series of other books: Macrolichens of the Northern Rocky Mountains (McCune and Goward 1995) A Colo r Guidebook to Common Rocky Mountain Lichens (St. Clair 1999) and A Rocky Mountain Lichen Primer (Corbridge and Weber 1998)

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12 In total, 19 species were positively identified (Appendix B ) during the photographic review. enotes an individual genus and species (eg., Acarospora contigua all encompassed species in relation to a site, including aspect, slope and quadrats. Then using data sheets and ticki ng off these counts, all species were tallied up per 1m 2 segment for a total of nine segments per quadrat. This was carried out for every upper and lower segment of each aspect as each site. Counts are approximations of species abundance, and the precision varied depending upon overlap of species on substrates. While multiple photographs were taken to improve visibility of counts, some photos in review had blurry edges or lighting that failed to lend a definitive identification of certain species. Only thos e species that could be identified through clear, focused, well lit photographic evidence were accounted for in this study. All numbers were then tallied per species, elevation, aspect, and site, before being entered into excel spreadsheets (see Table 3.2. 1 for total counts) Table 3.2.1 Totals of species accounted for by site, upper/lower, and aspect. VQS PIK GLA JSM Upper Total 4804 8277 8847 6420 Lower Total 3340 4883 8456 5380 Site Totals 8144 13160 17303 11800 Overall St udy Total 50407 Aspect Totals West 2472 2710 6160 3199 South 2864 3515 6861 3428 North 1590 2889 4282 3965 East 1218 4046 N/A 1208

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13 3.3 Statistical Analysis The data were analyzed in IBMs SPSS (IBMCorp. 2016) software for statis tical trends, as well as assessed with non parametric tests using the Simpson s Diversity Index (Simpson 1949) coefficient (Stohlgren et al. 1997) in order to conduct tests of species overlap, and a ranking of species abundance. A Generalized Li near Model (GLIM) (following Bolker et al. 2008) assuming a negative binomial distribution and log link function was used to compare the count data describing lichen species abu ndance across sites and aspects. S ite (four levels) and aspect (four levels) w ere treated as fixed effects. Following a statistically conservative approach, and due to a lack of a priori hypotheses about any expected site by aspect interactions, this factor was not included in the final GLIM. The generalized linear model with a nega tive binomial distribution was the approach recommended for the analysis of over dispersed count data (O'Hara and Kotze 2010, Bolker et al. 2008) Post hoc pairwise comparisons using the Bonferroni adjustment were made by site and aspect, and species count s were considered significantly different at p <0.05. Descriptive statistics of species abundance and averages across all sites, aspects, and distances to the summit (upper and lower) were also reported.

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14 CHAPTER IV R ESULTS A few initial general findi ngs stood out after the species were tallied for each of the sites (see Appendices I I IV ). Across all sites, 570 individual counts of lichen abundance were conducted with four site levels, four aspects (except for GLA), and two distances to summit (upper and lower), for a total of 30 quadrats, with 19 species observed within each First, sites totals for the three alpine sites (VQS, PIK and GLA) will be examined Then, the subalpine (JSM) sites totals and species counts a discussion of the Simpsons Diver sity Index and how it relates to each site follows. Next, species overlap across all four sites will be outlined, including a brief display of rank abundance. Finally, a review of overall statistical findings (bas ed on data obtained using SPSS) that were derived at the site level, aspect level and aspects within site is offered 4.1 VQS This site had the lowest overall total number of species at 8,144. The upper part of VQS had a total of 4,804 species, with the largest amount found on the southerly aspect (1,487). The most prevalent of species was Umbilicaria virginis at 1,380. Umbilicaria americana and Vulpicida pinastri both had zero presence. The lower portion of VQS had slightly lower numbers and had an appr oximate total of 3,340 species with the Southerly aspect once again exhibiting the largest numbers at (1,377). As with the upper aspect, U virginis (752) was the most prevalent species tallied, and V pinastri and Thamnolia subuliformis having zero presen ce at this distance from the summit.

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15 This site was fairly homogenous in cover between metamorphic rocks and grass in both the upper and lower portions examined and across all four aspects. This is reflected in the numbers in both aspect and relative dista nce to summit, with the upper portion having larger numbers due to there being slightly more rocks than grass (Fig. 4.1.1). Fig ure 4. 1 .1 Looking West to the summit from the lower East quadrat on VQS 4.2 PIK This site, lo cated north along the ridge from VQS had the second largest abundance of species at 13,160 and included all 19 species that have been identified in this study. This is in large part to the greater amount of rocks found at this site in comparison to the pre vious VQS site ( Fig. 4.2.1 ) The upper portion had an approximate total of 8,277 species with the

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16 highest proportion being on the Easterly (2,857). Diploschistes muscorum and T subuliformis soil based species, had zero presence in the upper portion of th is site and once again was dominated by U virginis (1,661). The lower portion had an approximate total of 4,883, with the largest total on the North (1,489). All 19 species were accounted for on the lower portion. Fig ure 4. 2 .1 view to the summit from the lower East quadrat at PIK 4.3 GLA This site was further north along the ridge and slightly East of the others, as it was situated along t he shoulder of Mt. Ida, (Fig. 4.3 .1). This site had the largest approximate abundan ce of species of all four sites with 17,303. Only three aspects were accounted for on

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17 this site due to the East aspect being on the edge of a steep cliff and thus inaccessible by the PI. The upper portion assessed had a total of 8,847 species accounted for with half of that being found on the Southerly (4,131). Both species of Vulpicida had zero presence, with U virginis dominating the rocks once again. The lower section of GLA had an approximate total of 8,456. All three aspects in the lower portion had t otals that were closer to one another than the upper with the Westerly aspect hosting the most (3,343). As in the upper portion and due to the abundance of rock cover, soil based species V pinastri V tilesii and T subuliformis had zero presence. U vir ginis once again dominated the rocks (3,575) closely followed by Rhizocarpon geographicum (3,504). The heavy presence of large, slabbed rock in the site ( Fig. 4.3.1) seemed to affect the high numbers of rock based species and lower numbers of soil based sp ecies. Figure 4. 3 .1 view to the North on GLA from the lower South quadrat

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18 4.4 JSM In contrast to the other three sites, JSM was situated west of the alpine ridge at a lower elevation, ( Fig. 4.4.1). This site had the third highest count in relation to the other three with an approximate total of 11,800. This site was very similar to VQS in that it was a balanced mix of rock and grass, with the exception of the East aspect. The East side appeared to have had some disturbance from some point in time, which left it with little rock and grass. This was a stark contrast to the other three aspects and was reflected in the numbers. Upper JSM had a total of 6,420 species with the North aspect ( Fig. 4.4 .2) accounting for 1,896. East upper was the lowest count of all four with 1,131. This trend was reflected in the lower portion of the East aspect where disturbance seemed high and only accounted for a total of 77 species, the lowest of any aspect from any site in this study. This seems to be in part of the difference in ground cover. Very few rocks, little grass, and various patches of soil dominated this portion of JSM. The lower part had a species total of 5,380 with the North aspect once again exhibiting the highest number at 2,069.

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19 Figure 4.4.1 Google Earth satellite imagery of the four study sites in RMNP Figure 4.4.2 the view South from the lower North aspect on JSM

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20 was implemented to test the lichen diversity within the datasets. Where n = total number of individuals of a particular species at a site (VQS, PIK, GLA, JSM) and N = total count of all species within the study sites, to characterize the ric hness, abundance, and evenness of lichen across sites (Arcadia 2013, Morris et al. 2014) The value of the index ranges from 0 to 1, with 0 having the lowest site diversity, and 1 the highest site diversity. The calculated high across all of the sites (range was 0.860 0.884), which was to be expected with 19 species found in the alpine and subalpine, which are common species for this region and these elevations. The lowest index was at VQS (0.86) and the highest at PIK ( 0.884) with GLA (0.864) and JSM (0.871) falling very closely in between. The probability of 87% that if one were to randomly choose two lichen individuals from the overall community assessed as the se sites, two different species would be picked in the sample, indicating a fairly high level of lichen diversity across sites. Site D= VQS 0.860 PIK 0.884 GLA 0.864 JSM 0.871

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21 Figure 4.5.1 A graphical represen from Table 4.5.1 per site location (VQS, PIK, GLA, JSM) oefficient parametric test to compare species overlap among sites, it was used for pairwise comparisons of observed lichen species identity overlap among sites and aspects, where A = the number of species (richness) found in both of the compared sites, B = the number of species found in the first site, but not present in the second site, and C = the number of species found in the second site, but not present in the first site (Stohlgren et al. 1997, Chong and Stohlgren 2007) to 0, then there is no overlap in species lists between sites, whereas if the coefficient is equal to 1.0, then there is complete overlap of all species across the two sites compared. Using this approach, this study first analyzed site diversity overlap using pairwise comparisons at the site level (see T able 4.6.2). Sites ranged from a low of 16 to a maximum of 19 species present (T able 4.6.1), and upon comparin 0.845 0.85 0.855 0.86 0.865 0.87 0.875 0.88 0.885 0.89 Sites VQS PIK GLA JSM

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22 0.947, indicating generally high species overlap across sites (with the 2 3 soil based lichen species commonly the ones absent, or in low abundance, across the sites except at PIK). was compared across plots that were within individual study sites (the subalpine site JSM and three alpine sites: VQS, PIK, GLA), and little variability was observed with respect to the Jaccard's coefficient in comparing overlap of within site richness bet ween aspects and proximity to the summit (upper and lower quadrats). Lastly, across sites, some notable differences aro se when comparing aspects. The northerly and westerly aspects exhibited large overlap in Jaccard's; however, when comparing the easterly and southerly aspects of the mean of the three alpine sites together (by pooling the upper and erlap was subst antially lower (T able aspects (mean of all three alpine sites), compared with these respective aspects in the and 0.74 comparing south. JSM, the subalpine site, had less exposed rock cover and vegetation broken up by patches of soil or loose rock. Table 4.6. 1 cies are represented per site VQS PIK GLA JSM 18 19 17 16

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23 Table 4.6. 2 Jaccard's pairwise comparisons Site Comparisons J = PIK 18/(18+0+1) 0.947 VQS JSM 16/(16+2+1) 0.842 GLA 18/(18+1+1) 0.9 PIK GLA 17/(17+2+ 0) 0.894 JSM 16/(16+3+0) 0.842 GLA JSM 16/(16+1+0) 0.941 Mean J = 0.896 Table 4.6. 3 Jaccard's East and South Alpine and Subalpine comparisons Jaccard's Alpine East = 0.615384615 compared to subalpine east Jaccard's Alpine South = 0.736842105 compared to subalpine south 4.7 Rank Abundance Rank abundance curves represent a way of plotting relative species abundance to visualize the biodiversity of species richness and evenness in a study. The cur ve displays the curves are a simple way for one to visualize whether species assessed are common or rare across sites. Data were obtained, per site, by taking the total for each species and dividing it by the site total. Based on the resulting quotient, each was assigned a rank (Table 4.7.1) and then graphed to show the resulting curves per site ( Fig 4.7.1). For all three Alpine sites (VQS, PIK, GLA) the top ranked species was Umbilicaria virginis while Acarospora contigua was top ranked at the subalpine site. Typically, there tend to be a few species within communities that exhibit dominance (maintain high abundance with respect to others), with

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24 the remainder of s pecies tending to be less common or rare, resulting in a dramatic drop and steep slope in species rank abundance curves. However, in this case with alpine and subalpine lichen, there exist a substantial number of species out of the 19 total which share a m oderately high abundance as indicated by the curve patterns observed in the Figure ( Fig. 4.7.1).

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25 Table 4.7.1 Rank abundance curves of species by site Rank VQS PIK GLA JSM 1 Umbilicaria virginis Umbilicaria virginis Umbilicaria virginis Acarospora c ontigua 2 Xanthoria elegans Rhizocarpon geographicum Rhizocarpon geographicum Lecidea atrobrunnea 3 Lecidea atrobrunnea Lecidea atrobrunnea Lecidea atrobrunnea Xanthoria elegans 4 Aspicilia caesiocinerea Acarospora contigua Umbilicaria decussata Aspicil ia caesiocinerea 5 Umbilicaria decussata Xanthoria elegans Acarospora contigua Rhizocarpon geographicum 6 Acarospora contigua Lecanora rupicola Lecidea tessellata Umbilicaria virginis 7 Lecanora rupicola Aspicilia caesiocinerea Lecanora rupicola Xanthop armelia cumberlandia 8 Xanthoparmelia wyomingica Umbilicaria decussata Aspicilia caesiocinerea Lecanora rupicola 9 Rhizocarpon geographicum Lecidea tessellata Xanthoria elegans Rhizoplaca chrysoleuca 10 Lecidea tessellata Rhizoplaca chrysoleuca Rhizopla ca chrysoleuca Xanthoparmelia wyomingica 11 Xanthoparmelia cumberlandia Pseudephebe minuscula Umbilicaria americana Xanthoparmelia chlorochroa 12 Rhizoplaca chrysoleuca Xanthoparmelia wyomingica Pseudephebe minuscula Umbilicaria americana 13 Xanthoparme lia chlorochroa Xanthoparmelia cumberlandia Xanthoparmelia wyomingica Pseudephebe minuscula 14 Pseudephebe minuscula Diploschistes muscorum Xanthoparmelia cumberlandia Lecidea tessellata 15 Diploschistes muscorum Xanthoparmelia chlorochroa Xanthoparmelia chlorochroa Umbilicaria decussata 16 Vulpicida tilesii Umbilicaria americana Diploschistes muscorum Diploschistes muscorum 17 Umbilicaria americana Vulpicida tilesii Thamnolia subuliformis Thamnolia subuliformis 18 Thamnolia subuliformis Thamnolia subu liformis Vulpicida pinastri Vulpicida pinastri 19 Vulpicida pinastri (0) Vulpicida pinastri Vulpicida tilesii Vulpicida tilesii

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26 Figure 4.7.1 Graph displaying the rank a bundance curve s of lichen species (1 19) ( proportional abundance with respect to rank abundance) per site 4.8 SPSS Final statistical analysis for the site and aspect species comparison s was carried out in IBMs SPSS statistical software (see Tables 4.8.1 through 4.8.5) With species site totals ranging from 0 to 3 575, these were n ot normal distributions and the raw count also exhibited overdispersion Trying to transform count data does not reliably improve analyses, necessitating a more conservative app roach to analysis (O'Hara and Kotze 2010) A 0.00 0.05 0.10 0.15 0.20 0.25 0.30 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Proportional Abundance Abundance Rank Rank Abundance Curves VQS PIK GLA JSM

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27 G eneral ized L i near M odel (GLIM) is a very flexible statistical model that works well with distributed dependent variables and allows for customization within the software itself. It also allows accommodation for empty cells (zero count values) and for customization to refine outputs within the study. For this study, species count was the dependent variable to be assessed against the four sites and their respective aspects for a total case processing value of 570 (19 species by 30 (summed from upper/lower aspects from all fours sites)). Indi vidual species counts contained a minimum of 0 and a maximum value of 1,037 (Table 4.7.1) A negative binomial probability distribution with log link function was assumed due to the fact that this count data exhibited a highly non normal distribution All assumptions were under the null hypothesis that no significant difference in species counts from site to site and when comparing aspects would be found This assumption was made because of the nature of the ecosystems contained in this study. As mentioned before, these are harsh environments, so diversity can be convoluted in comparison to lower elevation systems (Brodo et al. 2001b) The goodness of fit displays how well the values fit the distribution of the assumption of negative binomial with a log lin k (Tab le 4.7.3) Initially, p airwise comparisons were run putting each site up against the others. This was executed based on the dependent variable of count (species) first against site comparisons then against site aspects (Table 4.8.6). The Bonferroni c orrection was chosen as a more conservative approach over the l east significant difference (LSD) model as a correction for multiple comparis ons. This model, h owever, was not deemed appropriate because it resulted in everythin g showing significance. The pai rwise comparisons were then run with the Bonfer r oni correction, which corrects for post hoc multiple comparisons Using the Bonferroni correction for pairwise comparisons of differences in counts between sites or aspects, values below the p= 0 .05 level wer e

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28 considered significant With the site pairwise comparison s only the sites PIK and JSM, when compared were not statistically different ( p=0 .736). The total counts at these two sites were 13,160 and 11,800, respectively. The rest of the comparisons all s howed significant differences in lichen abundance between sites (Table 4.8.6 ) Finally, pairwise comparisons were conducted between aspects and only the South to East comparis ons showed any significance. Table 4.8 .1 This table display the overall data cou nt with its minimum, maximum, mean, and standard deviation Continuous Variable Information N Minimum Maximum Mean Std. Deviation Dependent Variable Count 570 0 1037 88.43 138.982 Table 4.8.2 Variables per site. N ote GLA has less due to the East a spect being excluded (Site Quadrats x 19 Species) Categorical Variable Information N Factor Site VQS 152 PIK 152 GLA 114 JSM 152 Total 570 Aspect West 152 South 152 North 152 East 114 Total 570

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29 Table 4.8 .3 This table di splays how well the data values fit the distribution with dependent variable count modeled against site and aspect Goodness of Fit a Value df Value/df Deviance 2011.923 563 3.574 Scaled Deviance 2011.923 563 Pearson Chi Square 1200.764 563 2.133 Scaled Pearson Chi Square 1200.764 563 Log Likelihood b 3082.607 Dependent Variable: Count Model: (Intercept), Site, Aspect a a. Information criteria are in smaller is better form. b. The full log likelihood function is displayed and used in compu ting information criteria. Table 4.8.4 Omnibus testing fitted model against intercept only models (site to aspect). Model effects tests the intercept model against site and aspect using negative binomial regression Omnibus Test a Tests of Model Effect s Likelihood Ratio Chi Square df Sig. Source Type III 90.971 6 0.000 Wald Chi Square df Sig. Dependent Variable: Count Model: (Intercept), Site, Aspect a (Intercept) 10509.880 1 0.000 a. Compares the fitted model against the intercept only model. Site 62.927 3 0.000 Aspect 16.959 3 0.001 Dependent Variable: Count Model: (Intercept), Site, Aspect

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30 Table 4.8.5 the reported mean and standard error for each site along with the 95% Wald confidence interval for upper and lower qua drats Estimated Marginal Means Site/Aspect Site Mean Std. Error 95% Wald Confidence Interval Lower Upper VQS 51.34 4.220 43.70 60.32 PIK 89.82 7.475 76.30 105.73 GLA 135.78 13.338 112.00 164.61 JSM 74.65 6.183 63.46 87.80 Aspect Mean Std. Erro r 95% Wald Confidence Interval Lower Upper West 90.57 7.417 77.14 106.34 South 104.08 8.504 88.68 122.15 North 80.39 6.616 68.41 94.46 East 61.68 6.223 50.61 75.17

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31 Table 4.8.6 Site to site comparisons using species c ount data as the dependent variable and the Bonfe r roni correction for multiple comparisons Pairwise Comparisons Site (I) Site Mean Difference (I J) Std. Error df Bonferroni Sig. 95% Wald Confidence Interval for Difference Lower Upper VQS PIK 38.4 7 a 8.629 1 0.000 61.24 15.71 GLA 84.44 a 13.911 1 0.000 121.14 47.74 JSM 23.30 a 7.478 1 0.011 43.03 3.57 PIK VQS 38.47 a 8.629 1 0.000 15.71 61.24 GLA 45.97 a 15.657 1 0.020 87.27 4.66 JSM 15.17 9.826 1 0.736 10.75 41.09 GLA VQS 84.44 a 1 3.911 1 0.000 47.74 121.14 PIK 45.97 a 15.657 1 0.020 4.66 87.27 JSM 61.14 a 14.494 1 0.000 22.90 99.38 JSM VQS 23.30 a 7.478 1 0.011 3.57 43.03 PIK 15.17 9.826 1 0.736 41.09 10.75 GLA 61.14 a 14.494 1 0.000 99.38 22.90 Pairwise comparisons of e stimated marginal means based on the original scale of dependent variable Count a. The mean difference is significant at the .05 level. Pairwise Comparisons Aspect (I) Aspect Mean Difference (I J) Std. Error df Bonferroni Sig. 95% Wald Confidence Interv al for Difference Lower Upper West South 13.50 11.255 1 1.000 43.20 16.19 North 10.19 9.937 1 1.000 16.03 36.40 East 28.89 a 9.796 1 0.019 3.05 54.74 South West 13.50 11.255 1 1.000 16.19 43.20 North 23.69 10.784 1 0.168 4.76 52.14 East 42.40 a 10.611 1 0.000 14.40 70.39 North West 10.19 9.937 1 1.000 36.40 16.03 South 23.69 10.784 1 0.168 52.14 4.76 East 18.71 9.103 1 0.239 5.31 42.72 East West 28.89 a 9.796 1 0.019 54.74 3.05 South 42.40 a 10.611 1 0.000 70.39 14.40 N orth 18.71 9.103 1 0.239 42.72 5.31 Pairwise comparisons of estimated marginal means based on the original scale of dependent variable Count

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32 CHAPTER V D ISCUSSION This study originally sought out to discover relationships of biological soil crusts (BSCs) in relation to high elevation sites in regards to (e.g., alpine and subalpine biomes). Once in situ, however, it was quickly discovered that the ground cover in these biomes contained few (if any) BSCs, but very abundant with lichen cover. Upon disc overing this, the research switched tracks, focusing on lichen exclusively. Within the sites themselves, the geology and land cover was fairly homogenous, especially in comparing the VQS and PIK site, with the only difference being PIK hosting a larger con gregation of metamorphic rocks. Overall, the geology remained similar, containing granite, gneiss, schist, and quartz outcrops The subalpine site held the least amount of rock cover for lichen habitability, and more particularly on the eastern side of JS M. This aspect hosted small areas of grass, but highly abundant soil patches, as if some process, such as a large herd of mammals, had disturbed the area. This aspect also contained the lowest count (77). No other aspect at any of the other sites had such drastically low lichen counts. The three alpine sites (VQS, PIK, GLA) ran along a ridge from North to South. Both VQS and GLA had their highest species counts on southerly aspects, with GLA containing the largest overall site total (17,303). When comparing the land cover between sites, GLA also had the most surface area of rocks than any other study site. The majority of lichen species at the GLA site were also known for colonizing rock faces. This is typical of above tree line ecosystems where mostly rocky ground remains the main landscape feature. Both Vulpicida pinastri/tilesii and Thamnolia subuliformis are well known soil based species, and Xanthoparmelia wyomingica is known to colonize both soil and rock. The high species count at GLA is most likely a result of these

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33 two environmental factors ( i.e. greater rock surface areas and the ability of certain species to thrive on rock specifically). The overall lichen species count totals seemed to lend more to differences among sites than some of the tests ra Diversity Index, the overall diversity yields variance across sites basically an 87% chance there was high species overlap a cross all four sites, which is to be expected on a smaller areal study with only 19 species of lichen identified. There was, however, a large overlap when comparing the east and south aspects among the means of alpine to subalpine. This was prevalent in th e pairwise aspect comparisons which were significant using the Bonferroni adjustment scores. Especially interesting, this analysis seemed to follow the higher totals for east and south aspects. In reviewing historical Google Earth imagery of the sites, a r ecurring snow bank was noticed, sometimes extending post winter along the eastern aspect of the ridgeline. The western aspect tends to experience a westerly wind pattern typical for that area of the Rocky Mountains, and this phenomenon, coupled with a clea red west side and higher snow accumulation on the east side, may be what could be affecting lichen totals. To determine if this is indeed a contributing factor to lichen numbers and colonization, further (snowfall) studies need to be conducted.

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34 CHAPTER VI C ONCLUSION High alpine ecosystems, such as those studied for this research, represent harsh environments for any organisms. Exposure in these locales remains a constant: from desiccating winds and UV rays, to drastic temperature fluctuations all inescapable on the bare earth with no shade. Resilience through biological adaptation represents a key component for species survival in such harsh extremes. Lichen, with their symbiotic relationship of algae/cyanobacteria housed inside a fungal body have managed to survive in all ecosystems across all continents on this planet. At its core, this study set out to discover spatial characteristics of such species. Though lichen dominated the area of study in between ground level vascular plants, this study te ased out a few relational findings among the 19 predominant species identified at the four study sites. wed all sites fell into a similar percentage (17 22%) across sites. The only differences were at VQS, which had (0 7%), the West side of JSM (22 26%) and the east aspect of GLA (31 37%), which was too steep to assess without the aid of climbing gear. These slopes were not steep enough to create a rock fall situation that may have affected the overall rock colonization of lichen present. In fact, most rocks were embedded in the ground at these sites. Due to the general evenness of slope across all four sites then, slope analyses were left out of this study. If proper safety gear could be utilized for assessing the east aspect of GLA, however, then slope could be worth investigating in a future follow up study. This could enhance findings in relation to the a spects and their influence on colonization.

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35 As the south and east facing aspects contained more total lichen than most of the others, other factors could be included in future studies, as a wide range of varying climatic factors (Grabherr, Gottfried and P auli 2000) would likely yield aspect relationships. For example, surrounding vegetation helps regulate moisture and soil temperature (Aalto, le Roux and Luoto 2013) and exposure to the sun and its path across the sky, will likely have an effect on those e lements. Typical westerly winds for the Rocky Mountain range would also likely play a part, as noted in the discussion and Google Earth images in Appendices F, when northern aspects remain cooler and moister than warmer southerly exposures (Selvakumar et al. 2009) Those climatic factors, in conjunction with, lichen habitat preferences, could infer aspect based populations based on solar and climatic data in relation to each of the four sites assessed. In addition to slope and aspect for possible future r esearch, the effects of carbon sequestration and nitrogen fixation could also be investigated. Most studies of this nature are conducted on soil based species only (De Deyn, Cornelissen and Bardgett 2008) and most often in controlled laboratory environmen ts (Yoshitake et al. 2010, Barger et al. 2006) where inputs into the soil can be measured. Inputs and variables associated with this aspect remain diverse: light, temperature, radiation, moisture, type of cyanobacteria or green algae, morphology, snow cove r, and permafrost, among others (Lange 2002, Lange and Green 1996, De Deyn et al. 2008) These prove difficult to measure due to the abundance of external influences (Lange and Green 2005, Lange, Reichenberger and Walz 1997) Though this study was not gear ed for such assessments, each of the above mentioned components could represent important topics for future research endeavors.

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36 In the end, while this study laid the basic foundation for future research endeavors related to alpine and sub alpine lichen sp atiality and identification, it also tested several statistical measures against somewhat large spatial parameters. This will allow future alpine and sub alpine lichen research to focus on more specific correlations, including climate related phenomena. Ov erall, this study will also inform other and further studies on lichen in alpine and sub alpine ecosystems of Rocky Mountain National Park specifically, and North America and other continents more generally.

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37 REFERENCES Aalto, J., P. C. le Roux & M. Luoto (2013) Vegetation Mediates Soil Temperature and Moisture in Arctic Alpine Environments. Arctic, Antarctic, and Alpine Research, 45 429 439. Allen, C. D. (2005) MICROMETEOROLOGY OF A SMOOTH AND RUGOSE BIOLOGICAL SOIL CRUST NEAR COON BLUFF, ARI ZONA. JOURNAL OF THE ARIZONA NEVADA ACADEMY OF SCIENCE, 38 21 28. --(2010) Biogeomorphology and Biological Soil Crusts: a symbiotic research relationship. Gomorphologie : relief, processus, environnement, Ames, S., K. Pischke, N. Schoenfuss, Z. Snob l, J. Soine, E. Weiher & T. Wellnitz (2012) Biogeographic patterns of lichens and trees on islands of the Boundary Waters Canoe Area Wilderness. Bios, 83 145 154. Arcadia, L. i. (2013) Lichen biogeography at the largest scales. The Lichenologist, 45 56 5 578. Ashton, I. 2011. Alpine vegetation composition, structure, and soils monitoring for Rocky Mountain National Park: 2010 Summary Report. In Natural Resource Data Series NPS/ROMN/NRDS 2011/148. Rocky Mountain Inventory and Monitoring Network: Nationa l Park Service. Barger, N. N., J. E. Herrick, J. Zee & J. Belnap (2006) Impacts of Biological Soil Crust Disturbance and Composition on C and N Loss from Water Erosion. Biogeochemistry, 77 247 263. Belnap, J. (2006) The potential roles of biological s oil crusts in dryland hydrologic cycles. Hydrological Processes, 20 3159 3178. Belnap, J., D. Eldridge, J. H. Kaltenecker, S. Leonard, R. Rosentreter & J. Williams. 2001. Biological Soil Crusts: Ecology and Management.

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38 Belnap, J. & O. L. Lange. 2003 a. Biological Soil Crusts of the Subalpine, Alpine, and Nival Areas in the Alps. In Biological Soil Crusts: Structure, Function, and Managment 67 72. Springer. --. 2003b. Biological Soil Crusts: Structure, Function, and Management New York: Springer Ver lag. Bjerke, J. W. (2008) Ice encapsulation protects rather than disturbs the freezing lichen. Plant Biology, 11 227 235. Bolker, B. M., M. E. Brooks, C. J. Clark, S. W. Geange, J. R. Poulsen, H. H. Stevens & J. S. S. White (2008) Generalized linear m ixed models: a practical guide for ecology and evolution. Trends in Ecology and Evolution, 24. Bonan, G. 2010. Ecological Climatology Concepts and Applications New York: Cambridge University Press. Bosch Roig, P., D. Barca, G. M. Crisci & C. Lalli (20 13) Lichens as bioindicators of atmospheric heavy metal deposition in Valencia, Spain. Journal of Atmospheric Chemistry, 70 373. Breen, K. & E. Lvesque (2008) The Influence of Biological Soil Crusts on Soil Characteristics along a High Arctic Glacier F oreland, Nunavut, Canada. Arctic, Antarctic, and Alpine Research, 40 287 297. Brodo, I. M., S. D. Sharnoff & S. Sharnoff. 2001a. Environmental Monitoring with Lichens. In Lichens of North America, ed. Y. U. P. N. H. London, 89 90. --. 2001b. Geographic Distributions in North America. In Lichens of North America 67. Yale University Press/New Haven and London. --. 2001c. Lichens of North America Yale University Press/New Haven and London. Chong, G. W. & T. J. Stohlgren (2007) Species area curves indi cate the importance of Ecological Indicators, 7 387 395. Corbridge, J. N. & W. A. Weber. 1998. A Rocky Mountain Lichen Primer Niwot, CO: University Press of Colorado.

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39 Cuculovic, A. A., M. S. Pavlovic, J. J. Savovic & D. S. Veselinovic (2014) DESORPTION OF METALS FROM CETRARIA ISLANDICA (L.) ACH. LICHEN USING SOLUTIONS SIMULATING ACID RAIN. Archives of Biological Sciences, 66 273 284. Dahl, E. (1954) Lichens. Botanical Review, 20 463 476. De Deyn, G. B., J. H. C. Cornelissen & R. D. Bardgett (2008) Plant functional traits and soil carbon sequestration in contrasting biomes. Ecology Letters, 11 516 531. Feuerer, T. & D. L. Hawksworth (2007) Biodiversity of lichens, including a world wide analysis Biodiversity and Conservation, 16 85 98. Gauslaa, Y. (2014) Rain, dew, and humid air as drivers of morphology, function and spatial distribution in epiphytic lichens. The Lichenologist, 46 1 16 Grabherr, G., M. Gottfried & H. Pauli (2000) GLORIA: A Global Observation Research Initiative in Alpine Environments. Mountain Research and Development, 20 190 191. Green, T. G. A. & O. L. Lange. 1995. Ecophysiology of Photosynthesis Springer Berl in Heidelberg. IBM SPSS Statistics for Windows, Version 24., IBM Corporation, Armonk, NY. Johnston, R. 1997. Introduction to Microbiotic Crusts. United States Department of Agriculture. Kar, S., A. C. Samal, J. P. Maity & S. C. Santra (2014) Diversit y of epiphytic lichens and their role in sequestration of atmospheric metals. International Journal of Environmental Science and Technology, 11 899 908.

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40 Karsten, U. & A. Holzinger (2012) Light, temperature, and desiccation effects on photosynthetic ac tivity, and drought induced ultrastructural changes in the green alga Klebsormidium dissectum (Streptophyta) from a high alpine soil crust. Microb Ecol, 63 51 63. Karsten, U., C. Ltz & A. Holzinger (2010) Ecophysiological Performance of the Aeroterrest rial Green Alga Klebsormidium Crenulatum (Charophyceae, Streptophyta) Isolated from an Alpine Soil Crust with an Emphasis on Desiccation Stress1. Journal of Phycology, 46 1187 1197. King, A. J., E. C. Farrer, K. N. Suding & S. K. Schmidt (2012) Co occur rence patterns of plants and soil bacteria in the high alpine subnival zone track environmental harshness. frontiers in MICROBIOLOGY, 3 347. Kularatne, K. I. A. & C. R. de Freitas (2013) Epiphytic lichens as biomonitors of airborne heavy metal pollution Environmental and Experimental Botany, 88 24 32. Lalley, J. S. & H. A. Viles (2007) Recovery of lichen dominated soil crusts in a hyper arid desert. Biodiversity and Conservation, 17 1 20. Lange, O. L. (2002) Photosynthetic productivity of the epil ithic lichen Lecanora muralis: Long term field monitoring of CO2 exchange and its physiological interpretationI. Dependence of photosynthesis on water content, light, temperature, and CO2 concentration from laboratory measurements. Flora, 197 233 249. La nge, O. L. & T. G. A. Green (1996) High Thallus Water Content Severely Limits Photosynthetic Carbon Gain of Central European Epilithic Lichens under Natural Conditions. Oecologia, 108 13 20. --(2005) Lichens Show That Fungi Can Acclimate Their Respirati on to Seasonal Changes in Temperature. Oecologia, 142 11 19. Lange, O. L., H. Reichenberger & H. Walz (1997) Continuous Monitoring of CO2 Exchange of Lichens in the Field: Short Term Enclosure with an Automatically Operating Cuvette. The Lichenologist, 29 259 274.

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41 McCune, B. & T. Goward. 1995. Macrolichens of the Northern Rocky Mountains Eureka, California: Mad Rivers Press, Inc. McIlroy de la Rosa, J. P., P. A. Warke & B. J. Smith (2012) Lichen induced biomodification of calcareous surfaces: Bio protection versus biodeterioration. Progress in Physical Geography, 37 325 351. Morris, K. E., T. Caruso, F. Buscot, M. Fischer, C. Hancock, T. S. Maier, T. Meiners, C. Muller, E. Obermaier, D. Prati, S. A. Socher, I. Sonnemann, N. Waschke, T. Wubet, S. Wurst & M. C. Rillig (2014) Choosing and using diversity indices: insights for ecological applications from the German Biodiversity Exploratories. Ecology and Evolution, 4 3514 3524. O'Hara, R. B. & D. J. Kotze (2010) Do not log transform count data. M ethods in Ecology and Evolution, 1 118 122. Pauli, H., M. Gottfried, D. Hohenwallner, K. Reiter, R. Casale & G. Grabherr. 2004. The Gloria Field Manuel Multi Summit Approach. ed. E. Union. Belgium. Peksa, O. & P. Skaloud (2011) Do photobionts influe nce the ecology of lichens? A casestudy of environmental preferences in symbiotic greenalga Asterochloris (Trebouxiophyceae). Molecular Ecology, 20 3936 3948. Pohl, M., D. Alig, C. Krner & C. Rixen (2009) Higher plant diversity enhances soil stability in disturbed alpine ecosystems. Plant and Soil, 324 91 102. Purvis, W. 2000. Lichens Smithsonian Institution Press. Raggio, J., T. G. A. Green & L. G. Sancho (2015) In situ monitoring of microclimate and metabolic activity in lichens from Antarctic e xtremes: a comparison between South Shetland Islands and the McMurdo Dry Valleys. Polar Biol, 39 113 122. Selvakumar, G., P. Joshi, P. K. Mishra, J. K. Bisht & H. S. Gupta (2009) Mountain Aspect Influences the Genetic Clustering of Psychrotolerant Phosp hate Solubilizing Pseudomonads in the Uttarakhand Himalayas. Current Microbiology, 59 432 438.

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42 Shukla, V. 2014. Selection of Biomonitoring Species. In Lichens to Biomonitor the Environment Springer India. Simpson, E. H. (1949) Measurement of Diversity Nature, 16 688. St. Clair, L. L. 1999. A Color Guidebook to Common Rocky Mountain Lichens Provo, Utah: M.L. Bean Life Science Museum Brigham Young University. Stephenson, S. L. 2010. The Kingdom Fungi Portland, OR: Timber Press, Inc. Stohlgren, T. J., G. W. Chong, M. A. Kalkhan & L. D. Schell (1997) RAPID ASSESSMENT OF PLANT DIVERSITY PATTERNS: A METHODOLOGY FOR LANDSCAPES. Environmental Monitoring and Assessment, 48 25 43. United States Department of Agriculture, N. R. C. S. 2007. Soil Survey of Rocky Mountain National Park, CO. ed. N. R. C. S. United States Department of Agriculture. Viles, H. A. (2008) Understanding Dryland Landscape Dynamics: Do Biological Crusts Hold the Key? Geography Compass 899 919. Walewski, J. 2007. Lichens of t he North Woods Duluth, MN: Kollath+Stensaas Publishing. Yoshitake, S., M. Uchida, H. Koizumi, H. Kanda & T. Nakatsubo (2010) Production of biological soil crusts in the early stage of primary succession on a high Arctic glacier foreland. New Phytol, 186 451 60. Zedda, L., A. Grngrft, M. Schultz, A. Peterson, A. Mills & G. Rambold (2011) Distribution patterns of soil lichens across the principal biomes of southern Africa. Journal of Arid Environments, 75 215 220. Zelikova, T. J., D. C. Housman, E. E. Grote, D. A. Neher & J. Belnap (2012) Warming and increased precipitation frequency on the Colorado Plateau: implications for biological soil crusts and soil processes. Plant and Soil, 355 265 282.

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43 APPENDIX A. LICHEN LIST FROM RMNP https://www.nps.g ov/romo/learn/nature/looking_for_lichens.htm Scientific Name Common Name Acarospora molybdina Acarospora schleicheri Allocetraria madreporiformis Arctoparmelia subcentrifuga Arthonia sp. Aspicilia caesiocinerea Aspicilia calcarea Aspi cilia sublapponica Bacidia subincompta Brodoa oroarctica Bryoria fuscescens Pale footed horsehair lichen Buellia leptocline Caloplaca exsecuta Caloplaca sinapisperma Caloplaca sp. 1 Caloplaca sp. 2 Caloplaca sp. 3 Caloplaca sp. 4 Caloplaca vitellinula Calvitimela armeniaca Candelariella sp. Candelariella terrigena Carbonea vorticosa Catapyrenium daedaleum Cetraria aculeata Cetraria ericetorum Iceland lichen Cetraria islandica Chaenotheca ferruginea Chromatochlamys muscorum octospora Cladonia cariosa Cladonia cenotea Powdered funnel lichen Cladonia cervicornis Ladder lichen Cladonia chlorophaea group Cladonia coccifera

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44 Cladonia coniocraea Common powderhorn Cladonia cornuta Bighorn liche n Cladonia cornuta groenlandica Cladonia deformis or sulphurina Cladonia didyma Southern soldiers Cladonia ecmocyna Frosted cladonia Cladonia ecmocyna intermedia Cladonia gracilis Cladonia macilenta bacillaris Cladonia macroceras Clado nia macrophyllodes Large leaved cladonia Cladonia phyllophora Cladonia sp. 1 Cladonia sp. 2 Cladonia sp. 3 Cladonia sp. 4 Cladonia sulphurina Cypehlium notarisii Dermatocarpon miniatum Leather lichen Dermatocarpon reticulatum Dipl oschistes muscorum Cowpie lichen Evernia divaricata Mountain oakmoss lichen Flavocetraria cucullata Flavocetraria nivalis Crinkled snow lichen Flavoparmelia caperata Flavoparmelia rutidota Hypogymnia bitteri Hypogymnia farinacea Icmadophi la ericetorum Imshaugia aleurites Ionaspis lacustris Lasallia papulosa Lasallia pensylvanica Lecanora allophana Lecanora chloropolia Lecanora epibryon Lecanora impudens Lecanora novomexicana Lecanora polytropa Lecanora rupi cola Lecanora sp.

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45 Lecanora thomsonii Lecidea atrobrunnea Lecidea promiscens Lecidea sp. Lecidea tessellata Tile lichen Lecidella scabra Leciophysma furfurascens Lepraria sp. Melanelia exasperatula Lustrous camouflage lichen Mel anelia glabroides Melanelia subolivacea Brown eyed camou f lage lichen Micarea tuberculata Miriquidica garovaglii Mycobilimbia berengeriana Nephroma bellum Nephroma parile Powdery kidney lichen Ochrolechia upsaliensis Parmelia fraudans Parmelia saxatilis Salted shield lichen Parmelia sulcata Hammered shield lichen Parmeliopsis ambigua Green starburst lichen Parmeliopsis hyperopta Gray starburst lichen Peltigera aphthosa Felt lichen Peltigera canina Dog lichen Peltigera collina Peltigera horizontalis polydactyla Peltigera malacea Veinless pelt Peltigera rufescens Field dog lichen Peltigera sp. Peltigera venosa Phaeophyscia constipata Physcia stellaris Star rosette lichen Physconia muscigena Placidium lachneum Placynthium nigrum Polyblastia cucurbitula Polychidium muscicola Porocyphus dispersus Porpidia superba Porpidia thomsonii Protoparmelia atriseda

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46 Protoparmelia badia Pseudephebe minuscula Pseudephebe pubescens? Psora luridel la Psora montana Psoroma hypnorum Green moss shingle Punctelia perreticulata Rhizocarpon disporum Rhizocarpon geographicum Yellow map lichen Rhizoplaca chrysoleuca Rhizoplaca melanophthalma Rinodina mniaraea Rinodina turfacea Tundra pepper spore lichen Schaereria cinereorufa Solorina crocea Orange chocolate chip lichen Staurothele clopimoides Stereocaulon sp. Stereocaulon tomentosum group Thamnolia subuliformis Toninia tristis Trapeliopsis granulosa Mottled disk li chen Umbilicaria americana Frosted rock tripe Umbilicaria decussata Umbilicaria deusta Umbilicaria hyperborea Blistered rock tripe Umbilicaria krascheninnikovii Salty rock tripe Umbilicaria lyngei Umbilicaria torrefacta Umbilicaria vellea A lpine rock tripe Umbilicaria virginis Usnea cavernosa Usnea lapponica Powdered beard lichen Vulpicida pinastri Powdered sunshine lichen Vulpicida tilesii Xanthoparmelia coloradoensis Colorado rock shield lichen Xanthoparmelia coloradoensis or cumberlandia Xanthoparmelia sp. Xanthoparmelia wyomingica Shingled rock shield lichen Xanthoria sorediata

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47 B. SPECIES TABLES Morphology Species Site Aspect Elevation Count Crustose Acarospora contigua VQS West Upper 76 Crustose Acarospora cont igua VQS West Lower 95 Crustose Acarospora contigua VQS South Upper 57 Crustose Acarospora contigua VQS South Lower 120 Crustose Acarospora contigua VQS North Upper 60 Crustose Acarospora contigua VQS North Lower 91 Crustose Acarospora contigua VQS Ea st Upper 65 Crustose Acarospora contigua VQS East Lower 57 Crustose Acarospora contigua PIK West Upper 97 Crustose Acarospora contigua PIK West Lower 101 Crustose Acarospora contigua PIK South Upper 127 Crustose Acarospora contigua PIK So uth Lower 52 Crustose Acarospora contigua PIK North Upper 85 Crustose Acarospora contigua PIK North Lower 141 Crustose Acarospora contigua PIK East Upper 278 Crustose Acarospora contigua PIK East Lower 275 Crustose Acarospora contigua GLA West Upper 131 Crustose Acarospora contigua GLA West Lower 198 Crustose Acarospora contigua GLA South Upper 201 Crustose Acarospora contigua GLA South Lower 236 Crustose Acarospora contigua GLA North Upper 157 Crustose Acarospora contigua GLA North L ower 210 Crustose Acarospora contigua GLA East Upper N/A Crustose Acarospora contigua GLA East Lower N/A Crustose Acarospora contigua JSM West Upper 258 Crustose Acarospora contigua JSM West Lower 323 Crustose Acarospora contigua JSM Sout h Upper 407 Crustose Acarospora contigua JSM South Lower 439 Crustose Acarospora contigua JSM North Upper 496 Crustose Acarospora contigua JSM North Lower 465 Crustose Acarospora contigua JSM East Upper 317 Crustose Acarospora contigua JSM East Lower 29

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48 Morphology Species Site Aspect Elevation Count Crustose Aspicilia caesiocinerea VQS West Upper 131 Crustose Aspicilia caesiocinerea VQS West Lower 75 Crustose Aspicilia caesiocinerea VQS South Upper 208 Crustose Aspicilia caesiocinerea VQS South L ower 140 Crustose Aspicilia caesiocinerea VQS North Upper 88 Crustose Aspicilia caesiocinerea VQS North Lower 57 Crustose Aspicilia caesiocinerea VQS East Upper 146 Crustose Aspicilia caesiocinerea VQS East Lower 88 Crustose Aspicilia cae siocinerea PIK West Upper 15 Crustose Aspicilia caesiocinerea PIK West Lower 9 Crustose Aspicilia caesiocinerea PIK South Upper 220 Crustose Aspicilia caesiocinerea PIK South Lower 29 Crustose Aspicilia caesiocinerea PIK North Upper 168 Crustose Aspic ilia caesiocinerea PIK North Lower 182 Crustose Aspicilia caesiocinerea PIK East Upper 227 Crustose Aspicilia caesiocinerea PIK East Lower 46 Crustose Aspicilia caesiocinerea GLA West Upper 42 Crustose Aspicilia caesiocinerea GLA West Lowe r 122 Crustose Aspicilia caesiocinerea GLA South Upper 140 Crustose Aspicilia caesiocinerea GLA South Lower 137 Crustose Aspicilia caesiocinerea GLA North Upper 44 Crustose Aspicilia caesiocinerea GLA North Lower 258 Crustose Aspicilia caesiocinerea G LA East Upper N/A Crustose Aspicilia caesiocinerea GLA East Lower N/A Crustose Aspicilia caesiocinerea JSM West Upper 226 Crustose Aspicilia caesiocinerea JSM West Lower 205 Crustose Aspicilia caesiocinerea JSM South Upper 350 Crustose As picilia caesiocinerea JSM South Lower 103 Crustose Aspicilia caesiocinerea JSM North Upper 155 Crustose Aspicilia caesiocinerea JSM North Lower 201 Crustose Aspicilia caesiocinerea JSM East Upper 126 Crustose Aspicilia caesiocinerea JSM East Lower 2

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49 Morphology Species Site Aspect Elevation Count Crustose Diploschistes muscorum VQS West Upper 0 Crustose Diploschistes muscorum VQS West Lower 0 Crustose Diploschistes muscorum VQS South Upper 3 Crustose Diploschistes muscorum VQS South Lower 17 Cru stose Diploschistes muscorum VQS North Upper 0 Crustose Diploschistes muscorum VQS North Lower 0 Crustose Diploschistes muscorum VQS East Upper 0 Crustose Diploschistes muscorum VQS East Lower 0 Crustose Diploschistes muscorum PIK West Upp er 0 Crustose Diploschistes muscorum PIK West Lower 22 Crustose Diploschistes muscorum PIK South Upper 0 Crustose Diploschistes muscorum PIK South Lower 6 Crustose Diploschistes muscorum PIK North Upper 0 Crustose Diploschistes muscorum PIK North Lowe r 0 Crustose Diploschistes muscorum PIK East Upper 0 Crustose Diploschistes muscorum PIK East Lower 1 Crustose Diploschistes muscorum GLA West Upper 1 Crustose Diploschistes muscorum GLA West Lower 17 Crustose Diploschistes muscorum GLA S outh Upper 3 Crustose Diploschistes muscorum GLA South Lower 1 Crustose Diploschistes muscorum GLA North Upper 1 Crustose Diploschistes muscorum GLA North Lower 4 Crustose Diploschistes muscorum GLA East Upper N/A Diploschistes muscorum GLA East Low er N/A Crustose Diploschistes muscorum JSM West Upper 18 Crustose Diploschistes muscorum JSM West Lower 5 Crustose Diploschistes muscorum JSM South Upper 5 Crustose Diploschistes muscorum JSM South Lower 0 Crustose Diploschistes muscorum JSM North Upper 9 Crustose Diploschistes muscorum JSM North Lower 0 Crustose Diploschistes muscorum JSM East Upper 0 Crustose Diploschistes muscorum JSM East Lower 0

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50 Morphology Species Site Aspect Elevation Count Crustose Lecanora rupicola VQS West Upper 23 Crustose Lecanora rupicola VQS West Lower 32 Crustose Lecanora rupicola VQS South Upper 42 Crustose Lecanora rupicola VQS South Lower 26 Crustose Lecanora rupicola VQS North Upper 44 Crustose Lecanora rupicola VQS North Lower 56 Crustose Le canora rupicola VQS East Upper 29 Crustose Lecanora rupicola VQS East Lower 30 Crustose Lecanora rupicola PIK West Upper 157 Crustose Lecanora rupicola PIK West Lower 91 Crustose Lecanora rupicola PIK South Upper 125 Crustose Lecanora rup icola PIK South Lower 61 Crustose Lecanora rupicola PIK North Upper 93 Crustose Lecanora rupicola PIK North Lower 84 Crustose Lecanora rupicola PIK East Upper 232 Crustose Lecanora rupicola PIK East Lower 113 Crustose Lecanora rupicola GL A West Upper 140 Crustose Lecanora rupicola GLA West Lower 185 Crustose Lecanora rupicola GLA South Upper 246 Crustose Lecanora rupicola GLA South Lower 144 Crustose Lecanora rupicola GLA North Upper 118 Crustose Lecanora rupicola GLA North Lower 133 Crustose Lecanora rupicola GLA East Upper N/A Crustose Lecanora rupicola GLA East Lower N/A Crustose Lecanora rupicola JSM West Upper 70 Crustose Lecanora rupicola JSM West Lower 62 Crustose Lecanora rupicola JSM South Upper 71 Crustose Lecanora rupicola JSM South Lower 10 Crustose Lecanora rupicola JSM North Upper 127 Crustose Lecanora rupicola JSM North Lower 139 Crustose Lecanora rupicola JSM East Upper 47 Crustose Lecanora rupicola JSM East Lower 5

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51 Morphology Species Site Aspe ct Elevation Count Crustose Lecidea atrobrunnea VQS West Upper 119 Crustose Lecidea atrobrunnea VQS West Lower 87 Crustose Lecidea atrobrunnea VQS South Upper 177 Crustose Lecidea atrobrunnea VQS South Lower 190 Crustose Lecidea atrobrunnea VQS North Upper 191 Crustose Lecidea atrobrunnea VQS North Lower 104 Crustose Lecidea atrobrunnea VQS East Upper 140 Crustose Lecidea atrobrunnea VQS East Lower 98 Crustose Lecidea atrobrunnea PIK West Upper 151 Crustose Lecidea atrobrunnea PIK Wes t Lower 191 Crustose Lecidea atrobrunnea PIK South Upper 314 Crustose Lecidea atrobrunnea PIK South Lower 124 Crustose Lecidea atrobrunnea PIK North Upper 150 Crustose Lecidea atrobrunnea PIK North Lower 151 Crustose Lecidea atrobrunnea PIK East Upper 356 Crustose Lecidea atrobrunnea PIK East Lower 178 Crustose Lecidea atrobrunnea GLA West Upper 489 Crustose Lecidea atrobrunnea GLA West Lower 505 Crustose Lecidea atrobrunnea GLA South Upper 627 Crustose Lecidea atrobrunnea GLA South L ower 517 Crustose Lecidea atrobrunnea GLA North Upper 349 Crustose Lecidea atrobrunnea GLA North Lower 411 Crustose Lecidea atrobrunnea GLA East Upper N/A Crustose Lecidea atrobrunnea GLA East Lower N/A Crustose Lecidea atrobrunnea JSM We st Upper 235 Crustose Lecidea atrobrunnea JSM West Lower 179 Crustose Lecidea atrobrunnea JSM South Upper 256 Crustose Lecidea atrobrunnea JSM South Lower 279 Crustose Lecidea atrobrunnea JSM North Upper 285 Crustose Lecidea atrobrunnea JSM North Lowe r 352 Crustose Lecidea atrobrunnea JSM East Upper 189 Crustose Lecidea atrobrunnea JSM East Lower 1

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52 Morphology Species Site Aspect Elevation Count Crustose Lecidea tessellata VQS West Upper 20 Crustose Lecidea tessellata VQS West Lower 35 Crustose Lecidea tessellata VQS South Upper 42 Crustose Lecidea tessellata VQS South Lower 44 Crustose Lecidea tessellata VQS North Upper 34 Crustose Lecidea tessellata VQS North Lower 17 Crustose Lecidea tessellata VQS East Upper 38 Crustose Lecidea tessella ta VQS East Lower 7 Crustose Lecidea tessellata PIK West Upper 29 Crustose Lecidea tessellata PIK West Lower 37 Crustose Lecidea tessellata PIK South Upper 94 Crustose Lecidea tessellata PIK South Lower 53 Crustose Lecidea tessellata PIK North Upper 93 Crustose Lecidea tessellata PIK North Lower 110 Crustose Lecidea tessellata PIK East Upper 133 Crustose Lecidea tessellata PIK East Lower 87 Crustose Lecidea tessellata GLA West Upper 102 Crustose Lecidea tessellata GLA Wes t Lower 187 Crustose Lecidea tessellata GLA South Upper 185 Crustose Lecidea tessellata GLA South Lower 162 Crustose Lecidea tessellata GLA North Upper 126 Crustose Lecidea tessellata GLA North Lower 223 Crustose Lecidea tessellata GLA East Upper N/A Crustose Lecidea tessellata GLA East Lower N/A Crustose Lecidea tessellata JSM West Upper 0 Crustose Lecidea tessellata JSM West Lower 0 Crustose Lecidea tessellata JSM South Upper 0 Crustose Lecidea tessellata JSM South Lower 0 Crustose Lecidea tessellata JSM North Upper 50 Crustose Lecidea tessellata JSM North Lower 0 Crustose Lecidea tessellata JSM East Upper 35 Crustose Lecidea tessellata JSM East Lower 6

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53 Morphology Species Site Aspect Elevation Count Fruticose Pseudephebe min uscula VQS West Upper 4 Fruticose Pseudephebe minuscula VQS West Lower 4 Fruticose Pseudephebe minuscula VQS South Upper 0 Fruticose Pseudephebe minuscula VQS South Lower 6 Fruticose Pseudephebe minuscula VQS North Upper 5 Fruticose Pseudephebe minusc ula VQS North Lower 1 Fruticose Pseudephebe minuscula VQS East Upper 0 Fruticose Pseudephebe minuscula VQS East Lower 0 Fruticose Pseudephebe minuscula PIK West Upper 23 Fruticose Pseudephebe minuscula PIK West Lower 22 Fruticose Pseudeph ebe minuscula PIK South Upper 78 Fruticose Pseudephebe minuscula PIK South Lower 11 Fruticose Pseudephebe minuscula PIK North Upper 6 Fruticose Pseudephebe minuscula PIK North Lower 24 Fruticose Pseudephebe minuscula PIK East Upper 82 Fruticose Pseude phebe minuscula PIK East Lower 0 Fruticose Pseudephebe minuscula GLA West Upper 40 Fruticose Pseudephebe minuscula GLA West Lower 18 Fruticose Pseudephebe minuscula GLA South Upper 50 Fruticose Pseudephebe minuscula GLA South Lower 37 Fru ticose Pseudephebe minuscula GLA North Upper 5 Fruticose Pseudephebe minuscula GLA North Lower 12 Fruticose Pseudephebe minuscula GLA East Upper N/A Fruticose Pseudephebe minuscula GLA East Lower N/A Fruticose Pseudephebe minuscula JSM Wes t Upper 35 Fruticose Pseudephebe minuscula JSM West Lower 3 Fruticose Pseudephebe minuscula JSM South Upper 7 Fruticose Pseudephebe minuscula JSM South Lower 0 Fruticose Pseudephebe minuscula JSM North Upper 0 Fruticose Pseudephebe minuscula JSM North Lower 5 Fruticose Pseudephebe minuscula JSM East Upper 42 Fruticose Pseudephebe minuscula JSM East Lower 0

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54 Morphology Species Site Aspect Elevation Count Crustose Rhizocarpon geographicum VQS West Upper 63 Crustose Rhizocarpon geographicum VQS Wes t Lower 3 Crustose Rhizocarpon geographicum VQS South Upper 28 Crustose Rhizocarpon geographicum VQS South Lower 27 Crustose Rhizocarpon geographicum VQS North Upper 63 Crustose Rhizocarpon geographicum VQS North Lower 27 Crustose Rhizocarpon geograph icum VQS East Upper 30 Crustose Rhizocarpon geographicum VQS East Lower 7 Crustose Rhizocarpon geographicum PIK West Upper 351 Crustose Rhizocarpon geographicum PIK West Lower 222 Crustose Rhizocarpon geographicum PIK South Upper 450 Crus tose Rhizocarpon geographicum PIK South Lower 48 Crustose Rhizocarpon geographicum PIK North Upper 124 Crustose Rhizocarpon geographicum PIK North Lower 201 Crustose Rhizocarpon geographicum PIK East Upper 648 Crustose Rhizocarpon geographicum PIK East Lower 316 Crustose Rhizocarpon geographicum GLA West Upper 718 Crustose Rhizocarpon geographicum GLA West Lower 579 Crustose Rhizocarpon geographicum GLA South Upper 869 Crustose Rhizocarpon geographicum GLA South Lower 633 Crustose Rhiz ocarpon geographicum GLA North Upper 307 Crustose Rhizocarpon geographicum GLA North Lower 398 Crustose Rhizocarpon geographicum GLA East Upper N/A Crustose Rhizocarpon geographicum GLA East Lower N/A Crustose Rhizocarpon geographicum JSM West Upper 125 Crustose Rhizocarpon geographicum JSM West Lower 118 Crustose Rhizocarpon geographicum JSM South Upper 185 Crustose Rhizocarpon geographicum JSM South Lower 5 Crustose Rhizocarpon geographicum JSM North Upper 174 Crustose Rhizocarpon ge ographicum JSM North Lower 426 Crustose Rhizocarpon geographicum JSM East Upper 232 Crustose Rhizocarpon geographicum JSM East Lower 31

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55 Morphology Species Site Aspect Elevation Count Foliose Rhizoplaca chrysoleuca VQS West Upper 4 Foliose Rhizoplac a chrysoleuca VQS West Lower 4 Foliose Rhizoplaca chrysoleuca VQS South Upper 3 Foliose Rhizoplaca chrysoleuca VQS South Lower 31 Foliose Rhizoplaca chrysoleuca VQS North Upper 41 Foliose Rhizoplaca chrysoleuca VQS North Lower 9 Foliose Rhizoplaca chr ysoleuca VQS East Upper 23 Foliose Rhizoplaca chrysoleuca VQS East Lower 6 Foliose Rhizoplaca chrysoleuca PIK West Upper 87 Foliose Rhizoplaca chrysoleuca PIK West Lower 46 Foliose Rhizoplaca chrysoleuca PIK South Upper 63 Foliose Rhizopl aca chrysoleuca PIK South Lower 52 Foliose Rhizoplaca chrysoleuca PIK North Upper 50 Foliose Rhizoplaca chrysoleuca PIK North Lower 54 Foliose Rhizoplaca chrysoleuca PIK East Upper 63 Foliose Rhizoplaca chrysoleuca PIK East Lower 22 Folio se Rhizoplaca chrysoleuca GLA West Upper 55 Foliose Rhizoplaca chrysoleuca GLA West Lower 114 Foliose Rhizoplaca chrysoleuca GLA South Upper 133 Foliose Rhizoplaca chrysoleuca GLA South Lower 105 Foliose Rhizoplaca chrysoleuca GLA North Upper 50 Folio se Rhizoplaca chrysoleuca GLA North Lower 11 Foliose Rhizoplaca chrysoleuca GLA East Upper N/A Foliose Rhizoplaca chrysoleuca GLA East Lower N/A Foliose Rhizoplaca chrysoleuca JSM West Upper 47 Foliose Rhizoplaca chrysoleuca JSM West Lower 92 Foliose Rhizoplaca chrysoleuca JSM South Upper 10 Foliose Rhizoplaca chrysoleuca JSM South Lower 82 Foliose Rhizoplaca chrysoleuca JSM North Upper 49 Foliose Rhizoplaca chrysoleuca JSM North Lower 69 Foliose Rhizoplaca chrysoleuca JSM East Upper 0 Foliose Rhizoplaca chrysoleuca JSM East Lower 0

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56 Morphology Species Site Aspect Elevation Count Fruiticose Thamnolia subuliformis VQS West Upper 0 Fruiticose Thamnolia subuliformis VQS West Lower 0 Fruiticose Thamnolia subuliformis VQS South Upper 1 Fruiticose Thamnolia subuliformis VQS South Lower 0 Fruiticose Thamnolia subuliformis VQS North Upper 0 Fruiticose Thamnolia subuliformis VQS North Lower 0 Fruiticose Thamnolia subuliformis VQS East Upper 0 Fruiticose Thamnolia subuliformis VQS East Lower 0 Fruiticose Thamnolia subuliformis PIK West Upper 0 Fruiticose Thamnolia subuliformis PIK West Lower 3 Fruiticose Thamnolia subuliformis PIK South Upper 0 Fruiticose Thamnolia subuliformis PIK South Lower 2 Fruiticose Thamnolia su buliformis PIK North Upper 0 Fruiticose Thamnolia subuliformis PIK North Lower 0 Fruiticose Thamnolia subuliformis PIK East Upper 0 Fruiticose Thamnolia subuliformis PIK East Lower 0 Fruiticose Thamnolia subuliformis GLA West Upper 0 Frui ticose Thamnolia subuliformis GLA West Lower 0 Fruiticose Thamnolia subuliformis GLA South Upper 1 Fruiticose Thamnolia subuliformis GLA South Lower 0 Fruiticose Thamnolia subuliformis GLA North Upper 0 Fruiticose Thamnolia subuliformis GLA North Lower 0 Fruiticose Thamnolia subuliformis GLA East Upper N/A Fruiticose Thamnolia subuliformis GLA East Lower N/A Fruiticose Thamnolia subuliformis JSM West Upper 0 Fruiticose Thamnolia subuliformis JSM West Lower 0 Fruiticose Thamnolia subuli formis JSM South Upper 0 Fruiticose Thamnolia subuliformis JSM South Lower 0 Fruiticose Thamnolia subuliformis JSM North Upper 0 Fruiticose Thamnolia subuliformis JSM North Lower 0 Fruiticose Thamnolia subuliformis JSM East Upper 0 Fruiticose Thamnoli a subuliformis JSM East Lower 0

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57 Morphology Species Site Aspect Elevation Count Foliose Umbilicaria americana VQS West Upper 0 Foliose Umbilicaria americana VQS West Lower 3 Foliose Umbilicaria americana VQS South Upper 0 Foliose Umbilicaria america na VQS South Lower 0 Foliose Umbilicaria americana VQS North Upper 0 Foliose Umbilicaria americana VQS North Lower 0 Foliose Umbilicaria americana VQS East Upper 0 Foliose Umbilicaria americana VQS East Lower 0 Foliose Umbilicaria america na PIK West Upper 11 Foliose Umbilicaria americana PIK West Lower 1 Foliose Umbilicaria americana PIK South Upper 3 Foliose Umbilicaria americana PIK South Lower 0 Foliose Umbilicaria americana PIK North Upper 0 Foliose Umbilicaria americana PIK North Lower 0 Foliose Umbilicaria americana PIK East Upper 5 Foliose Umbilicaria americana PIK East Lower 0 Foliose Umbilicaria americana GLA West Upper 12 Foliose Umbilicaria americana GLA West Lower 7 Foliose Umbilicaria americana GLA South Upper 71 Foliose Umbilicaria americana GLA South Lower 43 Foliose Umbilicaria americana GLA North Upper 32 Foliose Umbilicaria americana GLA North Lower 52 Foliose Umbilicaria americana GLA East Upper N/A Foliose Umbilicaria americana GLA East Lower N /A Foliose Umbilicaria americana JSM West Upper 47 Foliose Umbilicaria americana JSM West Lower 34 Foliose Umbilicaria americana JSM South Upper 8 Foliose Umbilicaria americana JSM South Lower 4 Foliose Umbilicaria americana JSM North Upp er 9 Foliose Umbilicaria americana JSM North Lower 24 Foliose Umbilicaria americana JSM East Upper 0 Foliose Umbilicaria americana JSM East Lower 0

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58 Morphology Species Site Aspect Elevation Count Foliose Umbilicaria decussata VQS West Upper 197 Fol iose Umbilicaria decussata VQS West Lower 63 Foliose Umbilicaria decussata VQS South Upper 169 Foliose Umbilicaria decussata VQS South Lower 202 Foliose Umbilicaria decussata VQS North Upper 108 Foliose Umbilicaria decussata VQS North Lower 21 Foliose Umbilicaria decussata VQS East Upper 3 Foliose Umbilicaria decussata VQS East Lower 8 Foliose Umbilicaria decussata PIK West Upper 115 Foliose Umbilicaria decussata PIK West Lower 82 Foliose Umbilicaria decussata PIK South Upper 185 Foli ose Umbilicaria decussata PIK South Lower 24 Foliose Umbilicaria decussata PIK North Upper 48 Foliose Umbilicaria decussata PIK North Lower 100 Foliose Umbilicaria decussata PIK East Upper 132 Foliose Umbilicaria decussata PIK East Lower 1 Foliose Umbilicaria decussata GLA West Upper 347 Foliose Umbilicaria decussata GLA West Lower 327 Foliose Umbilicaria decussata GLA South Upper 410 Foliose Umbilicaria decussata GLA South Lower 76 Foliose Umbilicaria decussata GLA North Upper 371 Fo liose Umbilicaria decussata GLA North Lower 150 Foliose Umbilicaria decussata GLA East Upper N/A Umbilicaria decussata GLA East Lower N/A Foliose Umbilicaria decussata JSM West Upper 24 Foliose Umbilicaria decussata JSM West Lower 5 Fol iose Umbilicaria decussata JSM South Upper 0 Foliose Umbilicaria decussata JSM South Lower 5 Foliose Umbilicaria decussata JSM North Upper 0 Foliose Umbilicaria decussata JSM North Lower 17 Foliose Umbilicaria decussata JSM East Upper 0 Foliose Umbili caria decussata JSM East Lower 0

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59 Morphology Species Site Aspect Elevation Count Foliose Umbilicaria virginis VQS West Upper 473 Foliose Umbilicaria virginis VQS West Lower 304 Foliose Umbilicaria virginis VQS South Upper 442 Foliose Umbilicaria vir ginis VQS South Lower 333 Foliose Umbilicaria virginis VQS North Upper 265 Foliose Umbilicaria virginis VQS North Lower 75 Foliose Umbilicaria virginis VQS East Upper 200 Foliose Umbilicaria virginis VQS East Lower 40 Foliose Umbilicaria virginis PIK West Upper 309 Foliose Umbilicaria virginis PIK West Lower 256 Foliose Umbilicaria virginis PIK South Upper 514 Foliose Umbilicaria virginis PIK South Lower 337 Foliose Umbilicaria virginis PIK North Upper 326 Foliose Umbilicaria virginis PIK North Lower 240 Foliose Umbilicaria virginis PIK East Upper 512 Foliose Umbilicaria virginis PIK East Lower 28 Foliose Umbilicaria virginis GLA West Upper 643 Foliose Umbilicaria virginis GLA West Lower 865 Foliose Umbilicaria virgin is GLA South Upper 1037 Foliose Umbilicaria virginis GLA South Lower 436 Foliose Umbilicaria virginis GLA North Upper 225 Foliose Umbilicaria virginis GLA North Lower 369 Foliose Umbilicaria virginis GLA East Upper N/A Foliose Umbilicaria virginis GLA East Lower N/A Foliose Umbilicaria virginis JSM West Upper 283 Foliose Umbilicaria virginis JSM West Lower 184 Foliose Umbilicaria virginis JSM South Upper 145 Foliose Umbilicaria virginis JSM South Lower 81 Foliose Umbilicaria virginis JSM North Upper 57 Foliose Umbilicaria virginis JSM North Lower 78 Foliose Umbilicaria virginis JSM East Upper 0 Foliose Umbilicaria virginis JSM East Lower 0

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60 Morphology Species Site Aspect Elevation Count Foliose Vulpicida pinastri VQS West Upper 0 Foliose Vulpicida pinastri VQS West Lower 0 Foliose Vulpicida pinastri VQS South Upper 0 Foliose Vulpicida pinastri VQS South Lower 0 Foliose Vulpicida pinastri VQS North Upper 0 Foliose Vulpicida pinastri VQS North Lower 0 Foliose Vulpicida pinast ri VQS East Upper 0 Foliose Vulpicida pinastri VQS East Lower 0 Foliose Vulpicida pinastri PIK West Upper 0 Foliose Vulpicida pinastri PIK West Lower 1 Foliose Vulpicida pinastri PIK South Upper 3 Foliose Vulpicida pinastri PIK South Lowe r 0 Foliose Vulpicida pinastri PIK North Upper 0 Foliose Vulpicida pinastri PIK North Lower 0 Foliose Vulpicida pinastri PIK East Upper 0 Foliose Vulpicida pinastri PIK East Lower 0 Foliose Vulpicida pinastri GLA West Upper 0 Foliose Vul picida pinastri GLA West Lower 0 Foliose Vulpicida pinastri GLA South Upper 0 Foliose Vulpicida pinastri GLA South Lower 0 Foliose Vulpicida pinastri GLA North Upper 0 Foliose Vulpicida pinastri GLA North Lower 0 Foliose Vulpicida pinastri GLA East Up per N/A Foliose Vulpicida pinastri GLA East Lower N/A Foliose Vulpicida pinastri JSM West Upper 0 Foliose Vulpicida pinastri JSM West Lower 0 Foliose Vulpicida pinastri JSM South Upper 0 Foliose Vulpicida pinastri JSM South Lower 0 Folio se Vulpicida pinastri JSM North Upper 0 Foliose Vulpicida pinastri JSM North Lower 0 Foliose Vulpicida pinastri JSM East Upper 0 Foliose Vulpicida pinastri JSM East Lower 0

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61 Morphology Species Site Aspect Elevation Count Foliose Vulpicida tilesii VQ S West Upper 0 Foliose Vulpicida tilesii VQS West Lower 0 Foliose Vulpicida tilesii VQS South Upper 0 Foliose Vulpicida tilesii VQS South Lower 0 Foliose Vulpicida tilesii VQS North Upper 0 Foliose Vulpicida tilesii VQS North Lower 2 Foliose Vulpicid a tilesii VQS East Upper 1 Foliose Vulpicida tilesii VQS East Lower 0 Foliose Vulpicida tilesii PIK West Upper 1 Foliose Vulpicida tilesii PIK West Lower 5 Foliose Vulpicida tilesii PIK South Upper 0 Foliose Vulpicida tilesii PIK South Lo wer 0 Foliose Vulpicida tilesii PIK North Upper 0 Foliose Vulpicida tilesii PIK North Lower 0 Foliose Vulpicida tilesii PIK East Upper 0 Foliose Vulpicida tilesii PIK East Lower 0 Foliose Vulpicida tilesii GLA West Upper 0 Foliose Vulpic ida tilesii GLA West Lower 0 Foliose Vulpicida tilesii GLA South Upper 0 Foliose Vulpicida tilesii GLA South Lower 0 Foliose Vulpicida tilesii GLA North Upper 0 Foliose Vulpicida tilesii GLA North Lower 0 Foliose Vulpicida tilesii GLA East Upper N/A Foliose Vulpicida tilesii GLA East Lower N/A Foliose Vulpicida tilesii JSM West Upper 0 Foliose Vulpicida tilesii JSM West Lower 0 Foliose Vulpicida tilesii JSM South Upper 0 Foliose Vulpicida tilesii JSM South Lower 0 Foliose Vulpicida t ilesii JSM North Upper 0 Foliose Vulpicida tilesii JSM North Lower 0 Foliose Vulpicida tilesii JSM East Upper 0 Foliose Vulpicida tilesii JSM East Lower 0

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62 Morphology Species Site Aspect Elevation Count Foliose Xanthoparmelia chlorochroa VQS West Up per 25 Foliose Xanthoparmelia chlorochroa VQS West Lower 1 Foliose Xanthoparmelia chlorochroa VQS South Upper 0 Foliose Xanthoparmelia chlorochroa VQS South Lower 4 Foliose Xanthoparmelia chlorochroa VQS North Upper 1 Foliose Xanthoparmelia chlorochro a VQS North Lower 9 Foliose Xanthoparmelia chlorochroa VQS East Upper 1 Foliose Xanthoparmelia chlorochroa VQS East Lower 0 Foliose Xanthoparmelia chlorochroa PIK West Upper 5 Foliose Xanthoparmelia chlorochroa PIK West Lower 0 Foliose Xa nthoparmelia chlorochroa PIK South Upper 0 Foliose Xanthoparmelia chlorochroa PIK South Lower 9 Foliose Xanthoparmelia chlorochroa PIK North Upper 2 Foliose Xanthoparmelia chlorochroa PIK North Lower 6 Foliose Xanthoparmelia chlorochroa PIK East Upper 1 Foliose Xanthoparmelia chlorochroa PIK East Lower 1 Foliose Xanthoparmelia chlorochroa GLA West Upper 3 Foliose Xanthoparmelia chlorochroa GLA West Lower 14 Foliose Xanthoparmelia chlorochroa GLA South Upper 23 Foliose Xanthoparmelia ch lorochroa GLA South Lower 17 Foliose Xanthoparmelia chlorochroa GLA North Upper 31 Foliose Xanthoparmelia chlorochroa GLA North Lower 2 Foliose Xanthoparmelia chlorochroa GLA East Upper N/A Foliose Xanthoparmelia chlorochroa GLA East Lower N/A Foliose Xanthoparmelia chlorochroa JSM West Upper 25 Foliose Xanthoparmelia chlorochroa JSM West Lower 29 Foliose Xanthoparmelia chlorochroa JSM South Upper 39 Foliose Xanthoparmelia chlorochroa JSM South Lower 0 Foliose Xanthoparmelia chlorochro a JSM North Upper 55 Foliose Xanthoparmelia chlorochroa JSM North Lower 21 Foliose Xanthoparmelia chlorochroa JSM East Upper 0 Foliose Xanthoparmelia chlorochroa JSM East Lower 0

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63 Morphology Species Site Aspect Elevation Count Foliose Xanthoparmelia cumberlandia VQS West Upper 35 Foliose Xanthoparmelia cumberlandia VQS West Lower 55 Foliose Xanthoparmelia cumberlandia VQS South Upper 41 Foliose Xanthoparmelia cumberlandia VQS South Lower 20 Foliose Xanthoparmelia cumberlandia VQS North Upper 11 Foliose Xanthoparmelia cumberlandia VQS North Lower 17 Foliose Xanthoparmelia cumberlandia VQS East Upper 14 Foliose Xanthoparmelia cumberlandia VQS East Lower 2 Foliose Xanthoparmelia cumberlandia PIK West Upper 0 Foliose Xanthoparmelia c umberlandia PIK West Lower 0 Foliose Xanthoparmelia cumberlandia PIK South Upper 1 Foliose Xanthoparmelia cumberlandia PIK South Lower 0 Foliose Xanthoparmelia cumberlandia PIK North Upper 5 Foliose Xanthoparmelia cumberlandia PIK North Lower 6 Folios e Xanthoparmelia cumberlandia PIK East Upper 97 Foliose Xanthoparmelia cumberlandia PIK East Lower 94 Foliose Xanthoparmelia cumberlandia GLA West Upper 31 Foliose Xanthoparmelia cumberlandia GLA West Lower 31 Foliose Xanthoparmelia cumber landia GLA South Upper 4 Foliose Xanthoparmelia cumberlandia GLA South Lower 23 Foliose Xanthoparmelia cumberlandia GLA North Upper 4 Foliose Xanthoparmelia cumberlandia GLA North Lower 3 Foliose Xanthoparmelia cumberlandia GLA East Upper N/A Foliose Xanthoparmelia cumberlandia GLA East Lower N/A Foliose Xanthoparmelia cumberlandia JSM West Upper 17 Foliose Xanthoparmelia cumberlandia JSM West Lower 14 Foliose Xanthoparmelia cumberlandia JSM South Upper 18 Foliose Xanthoparmelia cumber landia JSM South Lower 268 Foliose Xanthoparmelia cumberlandia JSM North Upper 253 Foliose Xanthoparmelia cumberlandia JSM North Lower 96 Foliose Xanthoparmelia cumberlandia JSM East Upper 43 Foliose Xanthoparmelia cumberlandia JSM East Lower 2

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64 Mor phology Species Site Aspect Elevation Count Foliose Xanthoparmelia wyomingica VQS West Upper 67 Foliose Xanthoparmelia wyomingica VQS West Lower 20 Foliose Xanthoparmelia wyomingica VQS South Upper 38 Foliose Xanthoparmelia wyomingica VQS South Lower 6 3 Foliose Xanthoparmelia wyomingica VQS North Upper 28 Foliose Xanthoparmelia wyomingica VQS North Lower 32 Foliose Xanthoparmelia wyomingica VQS East Upper 5 Foliose Xanthoparmelia wyomingica VQS East Lower 3 Foliose Xanthoparmelia wyomi ngica PIK West Upper 13 Foliose Xanthoparmelia wyomingica PIK West Lower 113 Foliose Xanthoparmelia wyomingica PIK South Upper 25 Foliose Xanthoparmelia wyomingica PIK South Lower 37 Foliose Xanthoparmelia wyomingica PIK North Upper 12 Foliose Xanthop armelia wyomingica PIK North Lower 10 Foliose Xanthoparmelia wyomingica PIK East Upper 12 Foliose Xanthoparmelia wyomingica PIK East Lower 0 Foliose Xanthoparmelia wyomingica GLA West Upper 6 Foliose Xanthoparmelia wyomingica GLA West Lowe r 53 Foliose Xanthoparmelia wyomingica GLA South Upper 13 Foliose Xanthoparmelia wyomingica GLA South Lower 31 Foliose Xanthoparmelia wyomingica GLA North Upper 16 Foliose Xanthoparmelia wyomingica GLA North Lower 2 Foliose Xanthoparmelia wyomingica G LA East Upper N/A Foliose Xanthoparmelia wyomingica GLA East Lower N/A Foliose Xanthoparmelia wyomingica JSM West Upper 67 Foliose Xanthoparmelia wyomingica JSM West Lower 80 Foliose Xanthoparmelia wyomingica JSM South Upper 13 Foliose Xa nthoparmelia wyomingica JSM South Lower 0 Foliose Xanthoparmelia wyomingica JSM North Upper 14 Foliose Xanthoparmelia wyomingica JSM North Lower 10 Foliose Xanthoparmelia wyomingica JSM East Upper 0 Foliose Xanthoparmelia wyomingica JSM East Lower 0

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65 Morphology Species Site Aspect Elevation Count Foliose Xanthoria elegans VQS West Upper 179 Foliose Xanthoria elegans VQS West Lower 275 Foliose Xanthoria elegans VQS South Upper 236 Foliose Xanthoria elegans VQS South Lower 154 Foliose Xanthoria el egans VQS North Upper 106 Foliose Xanthoria elegans VQS North Lower 27 Foliose Xanthoria elegans VQS East Upper 161 Foliose Xanthoria elegans VQS East Lower 16 Foliose Xanthoria elegans PIK West Upper 133 Foliose Xanthoria elegans PIK Wes t Lower 11 Foliose Xanthoria elegans PIK South Upper 321 Foliose Xanthoria elegans PIK South Lower 147 Foliose Xanthoria elegans PIK North Upper 238 Foliose Xanthoria elegans PIK North Lower 180 Foliose Xanthoria elegans PIK East Upper 79 Foliose Xan thoria elegans PIK East Lower 27 Foliose Xanthoria elegans GLA West Upper 57 Foliose Xanthoria elegans GLA West Lower 121 Foliose Xanthoria elegans GLA South Upper 118 Foliose Xanthoria elegans GLA South Lower 132 Foliose Xanthoria elegan s GLA North Upper 63 Foliose Xanthoria elegans GLA North Lower 145 Foliose Xanthoria elegans GLA East Upper N/A Foliose Xanthoria elegans GLA East Lower N/A Foliose Xanthoria elegans JSM West Upper 208 Foliose Xanthoria elegans JSM West L ower 181 Foliose Xanthoria elegans JSM South Upper 194 Foliose Xanthoria elegans JSM South Lower 444 Foliose Xanthoria elegans JSM North Upper 163 Foliose Xanthoria elegans JSM North Lower 166 Foliose Xanthoria elegans JSM East Upper 100 Foliose Xant horia elegans JSM East Lower 1

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66 C. SITE TABLES

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67 VQS

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68 PIK

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69 GLA

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70 JSM

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71 D. SITE TOTALS VQS Totals PIK Totals Acarospora contigua 621 Acarospora contigua 1156 Aspicilia caesiocinerea 933 Aspicilia caesiocinerea 896 Dip loschistes muscorum 20 Diploschistes muscorum 29 Lecanora rupicola 282 Lecanora rupicola 956 Lecidea atrobrunnea 1106 Lecidea atrobrunnea 1615 Lecidea tessellata 237 Lecidea tessellata 636 Pseudephebe minuscula 20 Pseudephebe minuscula 246 Rhizocarpon geographicum 248 Rhizocarpon geographicum 2360 Rhizoplaca chrysoleuca 121 Rhizoplaca chrysoleuca 437 Thamnolia subuliformis 1 Thamnolia subuliformis 5 Umbilicaria americana 3 Umbilicaria americana 20 Umbilicaria decussata 771 Umbilicaria decussata 687 Umbilicaria virginis 2132 Umbilicaria virginis 2522 Vulpicida pinastri 0 Vulpicida pinastri 4 Vulpicida tilesii 3 Vulpicida tilesii 6 Xanthoparmelia chlorochroa 41 Xanthoparmelia chlorochroa 24 Xanthoparmelia cumberlandia 195 Xanthoparmelia cumberlan dia 203 Xanthoparmelia wyomingica 256 Xanthoparmelia wyomingica 222 Xanthoria elegans 1154 Xanthoria elegans 1136 Site Total 8144 Site Total 13160

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72 GLA Totals JSM Totals Acarospora contigua 1133 Acarospora contigua 2734 Aspicili a caesiocinerea 743 Aspicilia caesiocinerea 1368 Diploschistes muscorum 27 Diploschistes muscorum 37 Lecanora rupicola 966 Lecanora rupicola 531 Lecidea atrobrunnea 2898 Lecidea atrobrunnea 1776 Lecidea tessellata 985 Lecidea tessellata 91 Pseudephebe minuscula 162 Pseudephebe minuscula 92 Rhizocarpon geographicum 3504 Rhizocarpon geographicum 1296 Rhizoplaca chrysoleuca 468 Rhizoplaca chrysoleuca 349 Thamnolia subuliformis 1 Thamnolia subuliformis 0 Umbilicaria americana 217 Umbilicaria americana 126 Umbilicaria decussata 1681 Umbilicaria decussata 51 Umbilicaria virginis 3575 Umbilicaria virginis 828 Vulpicida pinastri 0 Vulpicida pinastri 0 Vulpicida tilesii 0 Vulpicida tilesii 0 Xanthoparmelia chlorochroa 90 Xanthoparmelia chlorochroa 169 Xanthoparmelia cumberlandia 96 Xanthoparmelia cumberlandia 711 Xanthoparmelia wyomingica 121 Xanthoparmelia wyomingica 184 Xanthoria elegans 636 Xanthoria elegans 1457 Site Total 17303 Site Total 11800

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73 E. SPECIES PHOTOGRAPHS

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74 Acarospora contigua

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75 Aspicilia caesiocinerea

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76 Diploschistes muscorum

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77 Lecanora rupicola

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78 Lecidea atrobrunnea

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79 Lecidea tessellate

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80 Pseudephebe minuscula

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81 Rhizocarpon geographicum

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82 Rhizoplaca chrysoleuca

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83 Thamnolia subuliformis

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84 Umbilicaria Americana

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85 Umbilicaria decussate

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86 Umbilicaria virginis

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87 Vulpicida pinastri

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88 Vulpicida tilesii

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89 Xanthoparmelia chlorochroa

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90 Xanthoparmelia cumberlandia

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91 Xanthoparmelia wyomingica

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92 Xanthoria elegans

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93 F. GOOGLE EARTH HISTORICAL IMAGE RY Google Earth imagery of the study sites October 2013

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94 Google Earth imagery of the study sites July 2011

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95 G oogle Earth imagery of the study sites October 2005

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96 Google Earth imagery of the study sites September 1999

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97 Google Earth imagery of the study sites September 1990