Southeast Iowa Lichen Diversity and Elemental Analysis Amy Lynn Podaril Iowa State University

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2015 Southeast Iowa lichen diversity and elemental analysis Amy Lynn Podaril Iowa State University

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Southeast Iowa lichen diversity and elemental analysis

Amy Lynn Podaril

A thesis submitted to the graduate faculty

in partial fulfillment of the requirements for the degree of

MASTER OF SCIENCE

Major: Ecology and Evolutionary Biology

Program of Study Committee: James Colbert, Major Professor Leonor Leandro Brian Wilsey

Iowa State University

Ames, Iowa

2015

DEDICATION

This work is dedicated to all those that have encouraged and inspired me to study lichens.

iii

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS ...... v

ABSTRACT………………………………...... vi

CHAPTER 1 LICHEN BIOLOGY ...... 1

General ...... 1 Lichen Evolution ...... 2 Ecological/Economic Importance ...... 2 Lichens as Bioindicators ...... 4

CHAPTER 2 IOWA LICHEN HISTORY ...... 6

CHAPTER 3 INTRODUCTION OF STUDY SITES ...... 9

Wildcat Den State Park ...... 9 Big Sand Mound Nature Preserve...... 10 Starr’s Cave State Preserve ...... 10 Shimek State Forest ...... 11 The Sand Ridge- Private Land ...... 11

CHAPTER 4 LICHEN DIVERSITY IN SOUTHEAST IOWA……………………………………...... 13

Abstract ...... 13 Introduction ...... 13 Methods ...... 15 Results ...... 16 Discussion ...... 18 Acknowledgments...... 36 Literature cited ...... 37

CHAPTER 5 ELEMENTAL ANALYSIS OF IOWA PARMOTREMA SPECIES 41

Abstract ...... 41

Introduction ...... 41 Methods ...... 44 Results ...... 45 Discussion ...... 48 Acknowledgments...... 49 Literature Cited ...... 50

CHAPTER 6 SUMMARY AND CONCLUSIONS ...... 54

REFERENCES ...... 57

APPENDIX- IOWA LICHEN CHECKLIST 2015 ...... 62

ACKNOWLEDGEMENTS

I thank my committee chair, Jim Colbert, and my committee members, Leonor

Leandro, and Brian Wilsey for their guidance and support throughout the course of this research.

In addition, I thank my friends and family for their love and support throughout my studies. I also offer my appreciation to the many indiviuals who participated in field work as well as helping maintain the Iowa lichen database, without you this thesis would not have been possible.

ABSTRACT

Collection of 395 lichen specimens from five sites in southeast Iowa resulted in the identification of 150 species. Of these, 18 were species that had not been previously reported for the State of Iowa. Integration of these collections with previous records from a total of 30 herbaria allowed the estimate of 465 lichen species reported for Iowa. Lack of recent collection suggests that 33 of these species may be rare in, or extirpated from, the state.

Measurement of elemental concentrations was carried out in 8 populations of Parmotrema species from Shimek State Forest in southeast Iowa. The average elemental concentrations were 1.4 mg/g for phosphorus, 2.6 mg/g for potassium, 0.7 mg/g for iron, 2.8 mg/g for sulfur and 14.6 mg/g for nitrogen. Elemental concentrations were not significantly related to distance from forest edge.

CHAPTER I

LICHEN BIOLOGY

General

Lichens have been documented in the fossil record as far back as 600 million years ago (Yuan et al., 2005). This successful mutualistic relationship between fungi, algae and cyanobacteria continues to exist around the world encompassing over 17,600 known lichen species (Eriksson et al., 2004; Nelson et al., 2009). Lichens consist of two or three components. The fungal component of a lichen is called the mycobiont. The mycobiont of almost all lichens is a member of Phylum Ascomycota (Eriksson et al., 2004), although a small fraction of lichen species have a mycobiont belonging to Phylum Basidiomycota. The secondary and/or tertiary component is referred to as the photobiont (either a green algal species and/or cyanobacterial species). The mutualistic partners both benefit from the relationship, thriving under conditions that they may not normally survive in alone. The mycobiont provides its symbiont(s) with a stable environment, while the symbiont(s) provide the mycobiont with photosynthates.

The scientific name of a lichen is the scientific name of its mycobiont. Lichens are identified through distinctive anatomical, morphological, and chemical characteristics

(Brodo& Sharnoff, 2001). Still, identification of lichens is problematic in many instances, and studies involving DNA sequencing have provided more detailed taxonomic resolution

(Leavitt et al., 2011). As taxonomy changes and new species are discovered, it is important that all historical collections are updated to accurately determine species ranges and changes in species distribution over time. Some species that were once thought to be widely 2 distributed have been found to be separate species that are morphologically identical, but exist in distinguishable populations found in various locations throughout the world (Otálora,

2010). This could result in species being overlooked that are potential candidates for conservation efforts.

Lichen Evolution

Lichens are polyphyletic with approximately 10 independent lineages of lichens known (Nelson, et al., 2009). Evolutionary rates of symbionts have also been influenced by lichenization. Algal components have a much lower substitution rate as compared with the mycobionts. It is hypothesized that these low mutation rates in the algal component is controlled partially by the mycobiont, preventing genetic changes that would result in incompatibility between the two symbionts (Zoller & Lutzoni, 2003). This “preservation” of algal species within the thallus, could become important if, due to strong changes in the environment, free-living individuals are no longer available.

Fast rates of nucleotide substitutions in the mycobiont are more likely to occur in lichenizing lineages of fungi, as compared to non-mutualistic lineages (Lutzoni & Pagel,

1997). These quick spurts of mutation are hypothesized to lead to many different species of fungi, some losing the ability to lichenize. This highlights the importance of lichenization, as a mechanism for accelerated speciation in fungal evolution.

Ecological and Economic Importance

There are many ecological factors that could be negatively affected by decreases in lichen abundance or diversity. For example, cyanolichens play a substantial role in nitrogen fixation (Benner & Vitousek, 2007; Benner, 2011; Cambell et al., 2010). In the temperate oak savannah, nitrogen content of lichen litter on the forest floor is twice as high as that

3 found in oak leaves and twigs (Knops et al., 1996). Cyanolichens are also an important source of nitrogen for desert ecosystems, as well as preventing soil from major wind and water erosion (Evans, 1993, Belnap, 1995). Lichens also increase the amount of rainfall intercepted by the canopy in the temperate oak savanna by nearly 50% (Knops et al, 1996).

A wide variety of forest animals interact with lichens. Some birds and mammals, use macrolichens to build their nests (Hayward & Rosentreter, 1994). Some animal species consume lichens as a significant part of their diet. The most widely known example of this is caribou, which rely on the “reindeer lichen” for winter forage (Edwards, et al., 1960). There are many positive relationships between lichens and invertebrates as well. As lichen biomass increases the abundance of invertebrates (both terrestrial to aquatic) increases, along with a positive relationship between macrolichen species and spider abundance (Henderson &

Hackett, 1986; Stubbs, 1989; Gunnarson et al., 2004).

Lichenization has also played a key role in the production of secondary metabolites.

Both cyanobacteria and fungi produce secondary metabolites in the lichen thallus. Substances that are found in many lichens are predicted to act as a “sun screen” (along with other functions) providing protection from UV radiation for the photobionts (Gauslaa and Solhaug,

2001). Some lichen metabolites also serve as a protection mechanism from pathogenic microorganisms, animals, and surrounding vegetation (Lawrey, 1984). For example, in

South Wales, populations of certain grass species are restricted by an inhibitory chemical released by Peltigera canina (Pyatt, 1967). Accumulation of this chemical in the nearby soils diminishes root growth of native grasses, and has detrimental effects on germinating seedlings, preventing grasses from becoming invasive on the lichen territory. Overall, the production of secondary metabolites has been found in many fungal species, but is notably

4 common amongst lichenizing species and their descendants (Lutzoni et al., 2001).

Penicillium is an example of a non-lichenizing descendant, which produces a secondary metabolite called penicillin, a commonly used antibiotic.

Production of these secondary chemicals has led to lichens’ economic importance.

Some product ingredients include lichen extracts, such as perfumes and deodorants. For centuries lichens have been medically useful in treatments for cranial diseases, to cure jaundice, and treat stomach diseases (Huneck, 1999; Kirmizigül, 2003; Malhotra et al.,

2008). New compounds found in lichens show great potential in the area of medicine

(Malhotra et al., 2007). More recently extracts from three lichen species have been found to exhibit strong anticancer activity (Rankovic et al., 2011; Ghate et al., 2013). Substances found in lichens have also been used in research to degrade the prion protein in chronic wasting disease found in deer (Johnson et al., 2011; Bennett et al, 2012; Rodriguez et al.,

2012).

Lichens as Bioindicators

Lichens are used as bioindicators of air quality and pollutants from sulfur pollution to radioactive metals (Conti and Cecchetti, 2001). For example some lichen species are tolerant of high levels of nitrogen and other nutrients while other species are more sensitive. By observing whether sensitive species are present and healthy one can obtain a preliminary estimate of local nitrogen deposition levels. Similar observations can be made for species known to be sensitive to sulfur dioxide, heavy metals, and nitrogen.

Since lichens accumulate substances from the atmosphere the elemental concentration of the thallus can be measured. Although these concentrations are not direct measurements of atomospheric concentrations, they can be used to track sources of pollution

(such as livestock confinements or factories), or to compare concentration levels over time.

Tracking these levels was particularly useful after the incident in Chernobyl (Barci et al.,

1988; Seaward et al., 1988; Mihok et al., 1989; Livens et al, 1991; Sloof and Wolterbeck,

1992; Hoffmann et al., 1993; Triulzi et al., 1996; Sawidis et al., 1997). Not only were they able to validate the extent that the radiation traveled and reduction of radiation over time but also able to track radiation levels up the food chain from lichens to reindeer to the local people who consumed the reindeer (Jones et al., 1989, Rissanen and Rohala, 1989).

CHAPTER 2

IOWA LICHEN HISTORY

Lichen conservation has become an important topic as species (e.g., Lobaria pulmonaria) are declining around the world (Scheidegger, 2002; Zoller et al., 1999). There are currently no lichens listed on the State of Iowa Threatened or Endangered Species List

(Iowa Department of Natural Resources, 2014). The lack of lichens on the list is not reflective of their current status in the state, but instead a result of little available data.

Iowa has undergone severe landscape alterations since settlement by Europeans beginning in the early 1800s. In the 1830’s Iowa was covered in 28 million acres of tall grass prairie mixed with patches of wetlands and 7 million acres of forest (Johnson, 2000).

From1832-1875 Iowa’s forests dropped to fewer than 2.6 million acres (Cohen, 1992).

Virtually all of Iowa’s current forested land is second or third growth with very little or no old growth forest remaining in the state (Johnson, 2000). Plowing of greater than 99.9% of

Iowa’s 28 million acres of tall grass prairie followed beginning in the 1850’s, along with

95% of the prairie-wetland converted to agricultural use by 1970 (Cohen, 1992).

Knowledge of Iowa’s lichen diversity before and during these land use alterations is minimal. The earliest known lichen record for Iowa was Parmotrema crintium collected in

December of 1870 by C.E. Berry in Ames, Iowa (North American Lichen Consortium- WIS herbarium, 2014). Collection effort over the decades has been low and spotty (Figure 1).

Collections Made Per Year 800 700 600 500 400 300 200

Number Number of Collections 100

1870

1881

1888

1893

1898

1902

1908

1912

1916

1921

1925

1929

1934

1939

1948

1953

1957

1961

1965

1969

1973

1977

1981

1986

1991

1995

2001

2005

2009 2013 Year Figure 1 Number of lichens collected in Iowa each year. Collections from 2012-2013 include the study described in Chapter 4.

A large majority of Iowa lichen collections were made by Bruce Fink (1,353 specimens mostly collected between 1880-1903) and Clifford Wetmore (1,112 specimens collected between 1956-2006)

These historical collections were spatially restricted as most of Fink and Wetmore’s collections were concentrated in NE Iowa (especially Fayette County) (Figure 2). Although these collections are limited with respect to the locations sampled, they are invaluable to our knowledge of Iowa lichen diversity. Still, there are 70 of Iowa’s 99 counties with less than 50 total collections, and nine of Iowa’s counties that lack any collection effort to date.

The earliest published report of lichen diversity in the state (Bessey, 1884) noted the presence of 24 lichen species. Collection efforts in the 1970’s allowed Malone and Tiffany

(1978) to estimate that 263 lichen species were known from Iowa. The most recent effort

(Colbert, 2011) estimated a total of 448 lichen species reported for Iowa. Colbert (2011), considered species that hadn’t been collected in the state since 1960 to be “candidates for

Figure 2 Distribution map of Iowa lichen collections by county . Each point represents a single collection and the color indicates the year indicated in the legend. The points have been randomly placed in the county they were collected in in order to make them more visible (as many collections came from the same site). (Map produced in ArcGIS 10.2) rare or extirpated lichen species”. Only macrolichens were considered for this list, with a total of 42 species listed, although an additional 60 species of microlichens not collected since 1960 were also noted. Species that are rare or extirpated now, could clearly have been more common before these major land use changes occurred.

CHAPTER 3

INTRODUCTION TO STUDY SITES

A study of lichens in southeast Iowa is described below in Chapter 4. In this chapter, we will describe general background for each study site.

Wildcat Den State Park

Wildcat Den State Park covers 417 acres in Muscatine County Iowa. Exposed bedrock along Pine Creek is composed of limestone and dolomite from the Devonian period.

Other layers in the park include Pennsylvanian shale and sandstone. Glacial deposits from the most recent Illinoian glaciation are found in the park’s upland areas. Wind blown silt from more northern Wisconsinan glacial advance can also be found in the park. In the 1830’s the area was being logged and farmed. In 1927 the land was purchased by the State of Iowa and would later become Wildcat Den State Park. The forests of the park are dominated by oaks, honey locusts, sugar maple, and basswood. There are as many as 25 species of ferns that occur within the forest. The total vascular plant flora of the park exceeds 300 species.

(Anderson & Bunker 1997)

The city of Muscatine is located ten miles west of Wildcat Den, and Davenport is 15 miles east of the park. In 1848 Benjamin Nye built the Pine Creek Grist Mill used to grind buckwheat and rye into flour. This historic mill is currently being restored in the park.

Benjamin Nye was one of the first recorded settlers of Muscatine County (Anderson &

Bunker, 1997).

Big Sand Mound Nature Preserve

Big Sand Mound Nature Preserve is a 510 acre isolated sand prairie located in both

Louisa and Muscatine Counties along the Mississippi River. The preserve contains 352 native plants and 30 rare plants and animals (Wildlife Habitat Council, 2014). The site is well known for containing the endangered yellow mud turtle although its population is continuing to decline (Washburn, 2007). The Mid-American Energy Ecological Advisory Committee oversees management of the preserve. Prescribed burns have been practiced since 1986.

Twenty-nine acres of the northern section were burned in 2008. Modifications of Spring

Lake were installed to increase beneficial habitat for many species in the preserve (Wildlife

Habitat Council 2014).

Starr’s Cave State Preserve

Starr’s Cave State Preserve includes 184 acres of land about one mile outside of

Burlington. Limestone and dolomite bluffs stretch across the preserve. There are three caves located in the bluffs, but Starr’s Cave is the only naturally formed cave. The other two caves were manmade in the 1920’s in attempts to find zinc. Currently Starr’s cave has been closed to human access to prevent the spread of white nose in bats.

The land was originally a homestead to the Starr family in the 1860’s. Here they had a winery, cattle, sheep, and other livestock. The land was purchased by the DNR in the late

1970’s. The preserve is managed by the Des Moines County Conservation Board and is the hub of their environmental educational program. (Herzberg & Pearson, 2001)

Shimek State Forest

Shimek State Forest lies in both Lee and Van Buren Counties. The forest is named for Bohumil Shimek who was a University of Iowa faculty member and who collected numerous lichens that are currently in the University of Minnesota Herbarium. The forest was purchased in the 1930’s by the state along with some bordering abandoned farms creating one of the largest contiguous forests in Iowa. In our study we limited our research to only Lee County including portions of Donnellson and Farmington Units, and the entire Lick

Creek (which contains equestrian campground and trails), and Croton Units. The original function of Shimek State Forest was to serve as an example of forest management for Iowa’s citizens. Plantations of hardwoods and conifers were established and since 1972 have been used for commercial sales of sawtimber. The area contains limestone, shale, and sandstone bedrock. Area streams are slow running and are completely dry at times during the year. Oak, hickory, walnut, cottonwood, and elm are some of the common trees found in the forest.

About 1,000 acres consist of conifer plantations that include pines, spruce, and larch. The hardwood plantations include tulip poplar, swamp white oak, walnut, and black locust (Iowa

State Parks, 2014).

The Sand Ridge - Private land

The Sand Ridge is located on private land owned by Tom Sands in Fredonia, Iowa.

Much like Big Sand Mound this area consists of sand prairies and a high water table. This site however is much smaller (~30 acres) than all other sites and is completely surrounded by row crop (historically corn, soybeans, oats, and hay have been grown) and bordering an active railroad track on the south side. The entire property was purchased by the Sands

12 family in 1935, and portions of it had been previously farmed. The Sand Ridge itself has never been farmed due to its high sand content and inability to hold water.

CHAPTER 4 LICHEN DIVERSITY IN SOUTHEAST IOWA Submitted to Opuscula Philolichenum

Amy L. Podaril and James T. Colbert Ecology, Evolution, and Organismal Biology Department Iowa State University

Abstract

Exploration of the lichen diversity in southeast Iowa has been limited. Prior to this study, only

124 accessions (1.6%) of the total 7,644 known accessions of Iowa lichens were from four counties in southeast Iowa. After two years of field collection effort a total of 150 species were identified from 395 collected specimens, 18 of which are new records for the state. Only 29 of the species found had been previously documented in the southeast corner of the state.

Integration of these collecting efforts with previous efforts provides an estimate of 465 species of lichens reported for Iowa. Of these species, 33 macrolichen species are likely to be rare in, or extirpated (locally extinct) from, Iowa.

Introduction

Iowa has undergone severe landscape alterations since European settlement in the early 1800s. In the 1830’s, Iowa was covered in 28 million acres of tallgrass prairie mixed with patches of wetlands and 7 million acres of forest (Johnson, 2000). From1832-1875 Iowa’s forests dropped to fewer than 2.6 million acres (Cohen, 1992). Virtually all of Iowa’s current forested land is second or third growth with very little or no old growth forest remaining in the state (Johnson,

2000). Plowing of greater than 99.9% of Iowa’s 28 million acres of tallgrass prairie began in the

1850’s, along with 95% of the prairie-wetland converted to agricultural use by 1970 (Cohen,

1992). Knowledge of Iowa’s lichen diversity before and during these landuse alterations was

minimal. The earliest known lichen record for Iowa was Parmotrema crintium collected in

December of 1870 by C.E. Berry in Ames, Iowa (North American Lichen Consortium - WIS

herbarium, 2014). The earliest published report of lichen diversity in the state (Bessey, 1884)

noted the presence of 24 lichen species. The most recent effort (Colbert, 2011) estimated a total

of 448 lichen species reported for Iowa. Colbert (2011), considered species that had not been

collected in the state since 1960 to be “candidates for rare or extirpated lichen species”. Only

macrolichens were considered for this list, with a total of 42 species listed. Species that are rare

or extirpated now could clearly have been more common before these major landuse changes

occurred. To search for these rare species, as well as for species not previously identified in

Iowa, an under-collected portion of the state was selected to be surveyed. The southeast corner of

Iowa (Louisa, Lee, Des Moines, and Muscatine counties) has been poorly represented in

previous collections, with only 124 accessions (48 species) available before this study (Figure 1).

Figure 1 Map of Iowa with collection effort before study labeled in each county. Numbers in black font indicate the total number of accessions from that county. Numbers in red font indicate the number of accessions from before 1960.

Given the lack of knowledge about the lichen diversity in this region as well as its position in the

Mississippi River corridor – potentially providing a migration route for lichen species typically having a more southern distribution – southeast Iowa was the primary area of interest for this study.

Methods

Five sites were chosen for their accessibility and geographic features in an attempt to maximize the number of lichen species collected. These sites included Shimek State Forest (Lee County locations only), Big Sand Mound Preserve (Muscatine & Louisa County), Sand Ridge (Sand’s

Private Property- Louisa County), Starr’s Cave State Preserve (Des Moines County), and

Wildcat Den State Park (Muscatine County). Specimens were collected in winter and early spring months to avoid cover by vegetation. Habitats known to contain specific types of lichen communities were purposefully sought out and searched. The time spent in each location, as well as distance traveled while searching, was documented on each trip (Table 1). Georeference data were taken for each specimen collected along with habitat information. The collections were taken back to the lab where they were dried, frozen, identified (Harris & Ladd, 2014; Esslinger,

2003; Thomson, 2003; Orange et al., 2001; Śliwa, 2007; Smith et al., 2009; Sheard, 2010; Brodo,

2011; Lendemer, 2013) and deposited in the Ada Hayden Herbarium (ISC) at Iowa State

University. Thin layer chromatography (TLC) analysis was performed when necessary. Difficult specimens were sent to James Lendemer at the New York Botanical Garden for identification. A species list was created for each location to ensure that species that had already been collected at each site were not collected at the same site again. Species lists were created and compared throughout the study in attempts to find all available species at each site. Taxonomic nomenclature follows Esslinger, 2014.

Table 1 Collection sites and collecting effort.

Size Miles Hours Total New Record Site (Acres) Traveled Spent Species Species Shimek State 6,794 23.5 28.5 100 10 Forest Big Sand Mound 510 6.4 8 38 1 Preserve Sand Ridge 30 2.5 5 45 0 Wildcat Den State 413 7 10 49 5 Park Starr's Cave State 184 8.5 15 62 4 Preserve TOTAL 7,979 47.9 66.5 150 18

The Iowa Lichen Checklist (http://www.public.iastate.edu/~herbarium/IowaLichen/index.html) was developed using Colbert 2011, The North American Lichen Consortium online database, the

Iowa State Ada Hayden Herbarium lichen collection, and personal visits to Coe College, the

University of Nebraska-Lincoln, and the University of Minnesota. All taxonomic names were updated to follow Esslinger, 2014. The checklist was compared to the species found in southeast

Iowa. Any species that were not reported in the Iowa lichen species checklist, but were collected in the study were termed “new state record species”. The current likely extirpated list contains only macrolichens last collected before 1960. Species identification of specimens on the likely extirpated list were verified through a personal visit to the herbarium or receiving them on loan from the following herbaria: ASU, BRY, CINC, DUKE, FH, MICH, MSC, MONT, MU, NEB,

MIN, OMA, WIS, and NY. All likely extirpated ISC specimens have also been verified.

Results

Over the course of two years a total of 395 specimens were collected from the five sites in southeast Iowa. These collections corresponded to 150 lichen species (70% macrolichens), 119

17 of which had not been previously reported for southeast Iowa. Of these 150 species, 18 were new records for the State of Iowa (Table 2). Many of the species found in southeast Iowa (e.g.,

Physcia stellaris, Punctelia rudecta, etc.) are also common throughout other regions of the state.

Some species that are common in parts of Iowa (e.g., Teloschistes chrysophthalmus in southwest) Iowa are not common in southeast Iowa. Other species (e.g., Parmotrema perlatum) are relatively common in Shimek Forest but were not common in the other southeast Iowa collection sites. The following species are likely to be rare or uncommon for the state as they have not been found in locations other than southeast Iowa since 1960: Bacidia schweintzii (last collected in 1894 in northeast Iowa), Endocarpon diffractellum (last collected in 1905 in northeast Iowa), Eopyrenula intermedia (last collected in 1895 in northeast Iowa), Stictis urceolatum (last collected in 1895 in northeast Iowa), Cladonia scabriuscula (last collected in

1894 in northeast Iowa), and Usnea hirta (collected once in1895 in northeast Iowa). Only one of the new state record species has had no collections from surrounding states (Lecanora appalachensis). This species was recently described (Lendemer et al., 2013) and is a sterile crust lichen that requires TLC for identification. Therefore it is likely that it has been overlooked in these locations. One species collected in southeast Iowa had been listed on the Colbert (2011) likely extirpated list (Cladonia scabriuscula).

The accessions resulting from this study as well as previous accessions present in ISC and records from 29 additional herbaria have been compiled in the Iowa Lichen Database (Colbert,

2011; http://www.public.iastate.edu/~herbarium/IowaLichen/index.html), which currently includes about 8,000 accessions of lichens collected in Iowa. This total is likely to be an underestimate as some of the herbarium databases available on the Consortium of North

American Lichen Herbaria website (http://lichenportal.org/portal/) do not include records for all

18 of their holdings, and it is also likely that additional herbaria that have lichens collected from

Iowa are not part of the Consortium site. Still, the current Iowa Lichen Database provides our best understanding to date of both the historic and current lichen diversity of Iowa. Our current best estimate is that a total of 465 lichen species have been reported to occur in Iowa. Some of these species are very common and currently present in the state, such as Physcia stellaris with

262 accessions and collection dates ranging from 1882 to 2014, and Punctelia rudecta with 191 accessions and collection dates ranging from 1895 to 2014. Numerous other species (78) have only been documented on one occasion in the state of Iowa (e.g., Ionaspis alba). Ten of the 78 have not been collected since 1960, raising the question of whether they are rare in the state or simply difficult to find or identify. Such records also raise the issue of whether the specimen was correctly identified and are, perhaps, incorrectly included in Iowa’s lichen diversity. The current

Iowa Lichen Database includes a total of 94 species that have not been reported since 1960, 33 of which are macrolichens we consider to be likely rare or extirpated (Table 3). Specimens that have been personally viewed and are correctly identified are listed in the “verified” column.

Four of the likely rare or extirpated species have been reported to be acidophytic or sensitive to air pollution (Table 3).

Discussion

The collections made for this study add 18 species to the list of lichens reported for the State of

Iowa. Two of these species have a primarily southern distribution (Phlyctis boliviensis and

Lecanora appalachensis). Many of these new state record species were found in Shimek State

Forest (Table 1), which is not surprising as it is one of the largest contiguous forests in Iowa

(IDNR, 2014). Recently included records from other herbaria provide an additional 17 species compared with the most recent estimate (Colbert, 2011). Sixteen species were incorrectly

19 identified and removed from the list (e.g., Lecidea melaphanoides, Heppia despreauxii, and

Graphis eulectra). The work reported here increases the estimate of Iowa lichen diversity from

448 species (Colbert, 2011) to 465 species. This total is similar to three states that border Iowa:

Nebraska (411 species; Egan, et al, 2002), Wisconsin (726 species; Bennet, 2006), and

Minnesota (780 species, Bennet and Wetmore, 2004), suggesting that 465 species is a reasonable approximation of the diversity of lichens in Iowa, though further collection and cryptic species

(e.g., Lücking et al., 2014) could change that estimate. Many of these species (78) have only been reported once for the State of Iowa. This number may not accurately represent their abundance in the state as some these species are small and quite easy to overlook. In addition, nine of Iowa’s 99 counties have yet to have a single lichen species reported (Figure 1). Further investigation of those areas could reveal additional populations, additional species, or evidence regarding the status of potentially extirpated species.

Colbert (2011) listed 42 macrolichen species and noted 60 microlichen species as being potentially rare or extirpated based on the criterion of no reports in Iowa since prior to 1960.

This macrolichens list has been shortened to 33 after careful re-identification of herbarium specimens. One of the species collected in the present study (Cladonia scabriuscula) had been previously listed (Colbert, 2011) as being potentially rare or extirpated. Given that a single collection of C. scabriuscula was made in the current study, characterizing this species as “rare”, though not extirpated, in Iowa seems appropriate. None of the other 150 species collected in southeast Iowa were on the previous list of potentially rare or extirpated species, supporting the conclusion that the 33 species listed here are indeed likely to be rare or extirpated. Colbert

(2011) did extensive searching for Lobaria pulmonaria, but was unable to find it in the northeast portion state where it had been previously reported. This large and easily detected macrolichen

20 species was also not found in southeast Iowa. There is insufficient evidence available to make the claim that the 60 microlichen species not collected since 1960 are likely to be rare or extirpated as was noted by Colbert (2011).

The 33 macrolichen species that are likely to be rare or extirpated correspond to approximately

7% of the estimated total lichen diversity and approximately 14% of the macrolichen diversity in the state of Iowa. There are numerous potential explanations for rarity including the possibility that these species were rare in Iowa even prior to European settlement. The large number (22) of these macrolichen species last reported from Iowa prior to 1900, however, argues that at least some of these species have been extirpated from the state due to human activities. Dramatic changes in land use could certainly have led to extirpations, but other more subtle impacts could also play a role in some cases.

Some lichen species are known to be sensitive to airborne pollutants (Richardson, 1992). Some of these pollutants are deposited near farm and agricultural fields as dust or gases. Dust pollution can cause high concentrations of aluminum, iron, scandium, tin, and other elements (Nash,

2008), and today large hog farms are regulated to reduce levels of both air and water pollution caused by hydrogen sulfide and ammonia emissions (Sneeringer, 2010). Increases in atmospheric nitrogen deposition are a known global threat to the composition and function of ecosystems (Bobbink et al., 1998, 2010; Sala et al., 2000). The effects of anthropogenic nutrient inputs, especially nitrogen, on native plant vegetation is the topic of many studies (Gross et al.,

2005; Tilman and Lehman, 2001; Gratton & Denno, 2003). Results of these studies show both increases and decreases (depending on the species) in productivity, biodiversity, flowering, and tolerance to abiotic stresses (Phoenix et al., 2011; Galloway et al., 2004; Bobbink et al., 2010).

Increased nutrient level effects on lichen communities include: changes in lichen species

21 composition and cover, and changes in species distribution and colonization patterns, where species intolerant of eutrophic habitats are lost (Ruoss, 1999). There are 39 lichens on Iowa’s

Lichen Checklist that have been considered to be sensitive, acidophytic, or oligotrophic species in other studies (Geiser, 2014; Lawrey, 1993; Richardson, 1992; Wetmore, 1983, 1987, 1989,

1992, 1993a, 1993b, 1995). Some lichen species have conflicting sensitivity ratings in different geographic areas (Lawrey, 2014; USFS, 2014). Species such as Usnea hirta and Parmotrema crinitum are known to be both sensitive as well as oligotrophic species (USF, 2013; Wetmore,

1993a) and were unexpectedly found still persisting in the state. Perhaps Iowa’s climatic conditions, most importantly the fluctuations in temperature and moisture levels, aid in lichen tolerance of N deposition. High rains occurring while N deposition is at its peak can lessen the negative effects of atmospheric pollutants (Tretiach et al., 2012). Forest density and distance from agricultural land may also play a role in filtering out detrimental atmospheric levels of N and other pollutants.

Even if all 33 of the potentially extirpated macrolichens are, in fact, extirpated from the state,

Iowa still has a remarkable current lichen diversity of at least 400 species. This biological resource should be further explored giving attention to the diversity, distribution, and abundance of these organisms throughout the state. The 33 likely extirpated macrolichen species (Table 3) should be considered for listing as “special concern species” by the State of Iowa so that appropriate conservation measures could be taken for any populations that may be discovered in the future. Consideration should also be given to reintroduction of extirpated lichen species for which suitable habitat can be identified.

Table 2 Lichen species collected in southeast Iowa. New state record species are highlighted in green. The “#” column represents the total number of known accessions for each species (including collections outside of southeast Iowa). Rare/uncommon species are indicated with “*” in the “#” column. Sensitivity abbreviations are the following: “S” means sensitive to sulfur and other air pollutants, “I” indicates species with intermediate sulfur and other air pollutant sensitivity, “T” indicates sulfur and other air pollutant tolerance, “S-I” indicates species between sensitive and intermediate, “N” indicates nitrophytic species, “A” indicates acidophytic species, “E” indicates eutrophic species, “O” indicates oligotrophic species, “M” indicates mesotrophic species. All sensitivity ratings that were contradictory are not listed. States adjacent to Iowa that have collections of new record or rare/uncommon species are listed.

Adjacent Genus Species Author Synonyms # Sensitivity States

(Ach.) R. C. Arthopyrenia Acrocordia cavata 8 Harris cavata Arthopyrenia Acrocordia conoidea (Fr.) Körber 2 conoidea

(Pers.) Ertz & Tehler (Ertz Opegrapha Alyxoria varia 87 & Tehler diaphora, O. varia 2011)

Rinodina dakotensis, R. (H. Magn.) P. finkii, R. Amandinea dakotensis May & inaequalis, R. 30 Sheard pennsylvanica, R. subplumbea, R. subpyriniformis

(Hoffm.) Buellia punctata, Amandinea punctata Coppins & B. myriocarpa, B. 65 N Scheid. pullata

Arthopyrenia biformis, A. parvula, A. (Borrer) R. C. conformis auct. N. Anisomeridium biforme 4 Harris Am., Ditremis biformis, Trimmatothele umbellulariae

Anisomeridium nyssaegenum, (Ellis & Ditremis Anisomeridium polypori Everh.) M.E. 12 nyssigena, Barr Arthopyrenia willeyana

(Pers.) A. Arthonia atra 2 I Schneider

23 Table 2 continued

Arthonia patellulata Nyl. 9

Arthonia radiata (Pers.) Ach. 49 I, A

(Flotow) A. Arthonia Arthothelium spectabile 34 Massal. spectabilis

(Hoffm.) Caloplaca Athallia holocarpa Arup, Frödén 44 N holocarpa & Søchting

B. trachona, Bilimbia (Körber) pammellii, Bacidia coprodes 47 Lettau Bilimbia granosa, Bilimbia trachona auct.

(Hoffm.) A. Bacidia rubella 12 I Massal.

(Fr. ex E. Bacidia schweinitzii Michener) A. Biatora leucampyx 3* Schneider

(Fr.) A. Bacidia suffusa Biatora suffusa 26 Schneider

Bacidina egenula (Nyl.) Vězda Bacidia egenula 10

Bacidina inundata (Fr.) Vězda Bacidia inundata 41

(DC.) Verrucaria Bagliettoa calciseda Gueidan & 3 calciseda Cl. Roux

IL, MI,MN, Biatora vernalis (L.) Fr Lecidea vernalis 1 WI

24 Table 2 continued

Bilimbia Bacidia sabuletorum, B. hypnophila, (Schreber) sabuletorum Mycobilimbia 40 Arnold sabuletorum, Myxobilimbia sabuletorum

(Ehrh. ex Caloplaca cerina Hedw.) Th. C. ulmorum 144 S-I Fr.

Caloplaca microphyllina (Tuck.) Hasse 73

Vondrák & Caloplaca soralifera 6 Hrouzek Coppins & P. Caloplaca ulcerosa 11 James

(Dickson) Candelaria concolor 200 E, N Stein

(Fr.) Mull. Candelaria fibrosa 66 Arg.

(Hoffm.) Candelariella aurella 7 N Zahlbr.

R.C. Harris & Candelariella efflorescens 9 W.R. Buck

Borrer) A. Catillaria chalybeia 2 Massal.

Catillaria glauconigrans (Tuck.) Hasse 1 WI, SD

Bacidia declinis, Catillaria nigroclavata (Nyl.) Schuler Bilimbia declinis, 4 Lecidea declinis

25 Table 2 continued

Chrysothrix Allarthonia caesia, (Flotow) Arthonia caesia 93 Körber lecideella, Arthonia caesia

Cladonia apodocarpa Robbins 9

Cladonia caespiticia (Pers.) Flörke 5

(Ach.) Cladonia cariosa 25 Sprengel

(Flörke ex Cladonia chlorophaea Sommerf.) 43 S, I, E Sprengel

(Flörke) Cladonia coniocraea 26 I, S Sprengel WI, IL,SD, Cladonia crispata Ach. 2 MN

Cladonia cristatella Tuck. 32 I

Cladonia cryptochlorophaea Asahina 6

(Flörke) Cladonia decorticata 7 Sprengel

Cladonia fimbriata (L.) Fr. 14 S-I, S, E

(Hudson) Cladonia furcata 73 I, O, T Schrader

Cladonia macilenta Hoffm. 23 S, E

(Hoffm.) Cladonia parasitica 35 Hoffm.

(With.) J. R. C. mitrula, C. Cladonia peziziformis 55 Laundon capitata

Cladonia pyxidata (L.) Hoffm. 44 M

Cladonia rei Schaerer 17

26 Table 2 continued

Cladonia robbinsii A. Evans 10

MI, MN, Cladonia scabriuscula (Delise) Nyl. 2* SD, WI,

Cladonia squamosa Hoffm. 21 I, E

(Ach.) Grogn Cladonia strepsilis 1 MO, WI ot C. clavulifera, C. WI, IL, NE, Cladonia subcariosa Nyl. 8 polycarpoides MO (L.) F. H. Cladina Cladonia subtenuis 12 Wigg rangiferina (L.) F. H. Cladonia subulata 2 MN, IL Wigg (Flörke) Fr. (Ahti 2000, Cladonia symphycarpa 18 Hansen & Ahti 2011)

(Ach.) Dimerella pineti, WI, SD, Coenogonium pineti Lücking & D. diluta, 1 MO, MN, Lumbsch Microphiale diluta IL

Collema subflaccidum Degel. 12

Synechoblastus Collema texanum Tuck. texanus, S. 1 laciniatus

Cyphelium tigillare (Ach.) Ach. 26

(With.) J. R. Dermatocarpon luridum 5 Laundon

Dermatocarpon miniatum (L.) W. Mann 107

(Scop.) R. Diploschistes muscorum 12 Sant.

(Nyl.) Staurothele Endocarpon diffractellum Gueidan & diffractella, 7*

Cl. Roux Verrucaria

27 Table 2 continued diffractella

Endocarpon pusillum Hedwig 78

Parmelia (Gyelnik & baltimorensis, Flavoparmelia baltimorensis 5 Fóriss) Hale Pseudoparmelia baltimorensis

Parmelia caperata, P. cylisphora, P. flavicans, P. Flavoparmelia caperata (L.) Hale herreana, P. 123 I, E, N negativa, Pseudoparmelia caperata

(Hoffm.) Th. Flavoplaca citrina Caloplaca citrina 29 N Fr.

Graphis scripta (L.) Ach. 159 I

(Hudson) Søchting, Caloplaca Gyalolechia flavorubescens 107 Frödén & flavorubescens Arup

(Wulfen) Søchting, Caloplaca Gyalolechia flavovirescens 71 Frödén & flavovirescens Arup

Physcia adglutinata, P. (Flörke) H. elaeina, Hyperphyscia adglutinata Mayrhofer & 53 I, A Physciopsis Poelt adglutinata, P. elaeina

Physcia syncolla, (Tuck. ex Hyperphyscia syncolla Physciopsis 133 Nyl.) Kalb syncolla

Lecania cyrtella (Ach.) Th. Fr 1 A SD, MN

(Zahlbr.) MI, MO, Lecania croatica 1 Kotlov WI, IL

Table 2 continued Lecania erysibe (Ach.) Mudd 6

Lendemer & Lecanora appalachensis 1 R.C. Harris

Lecanora chlarotera Nyl. L. rugosella 9 I

(Pers.) Lecanora dispersa 40 T, N Sommerf.

(Sprengel) Lecanora strobilina 17 Kieffer

Lecanora symmicta (Ach.) Ach. Lecidea symmicta 21

Lecanora thysanophora Harris 14

Crocynia aliciae, (B. de Lesd.) Lepraria finkii C. americana, L. 69 A R. C. Harris lobificans

Parmelina aurulenta, (Tuck.) Elix Myelochroa aurulenta Parmelia 77 & Hale aurulenta, P. silvestris

Parmelina galbina, (Ach.) Elix & Parmelia galbina, Myelochroa galbina 31 Hale P. subquercifolia, P. sulphurosa

Opegrapha herbarum Mont. 4

Opegrapha vulgata Ach. 2 I

Parmelia Parmotrema austrosinense (Zahlbr.) Hale 1 WI, MO, IL austrosinensis Parmelia crinita, (Ach.) M. Parmotrema crinitum Parmelia 20 S,O Choisy proboscidea

Parmelia hypotropa, Parmelia Parmotrema hypotropum (Nyl.) Hale hypotropa var. 10 T sorediata , P. cetrata var. hypotropoides

Table 2 continued 29

Parmotrema Parmelia perlata, (Hudson) M. P. trichotera, perlatum 5 A Choisy Parmotrema chinense

Parmelia (Taylor) M. Parmotrema reticulatum reticulata, Rimelia 17 S Choisy reticulata

Peltigera canina (L.) Willd. 82 T

(Florke ex Peltigera praetextata Sommerf.) 33 S Zopf

(I. M. Lamb) WI, MN, Pertusaria macounii 1 Dibben MO

Pertusaria pustulata (Ach.) Duby 41

Pertusaria rubefacta Erichsen 1

Phaeophyscia adiastola (Essl.) Essl. Physcia adiastola 57

(Hoffm.) Physcia ciliata, P. Phaeophyscia ciliata 139 Moberg obscura auct.

Phaeophyscia (Mereschk.) Phaeophyscia hirsuta cernohorskyi, 57 Essl. Physcia hirsuta

Phaeophyscia hirtella Essl. 20

Physcia hispidula, Phaeophyscia hispidula (Ach.) Essl. 19 P. setosa

Physcia (Necker) Phaeophyscia orbicularis orbicularis, P. 40 I, E, N Moberg virella

Phaeophyscia pusilloides (Zahlbr.) Essl. Physcia pusilloides 29

Physcia orbicularis f. Phaeophyscia rubropulchra (Degel.) Essl. rubropulchra, P. 64 N endochrysea, P. rubropulchra

30 Table 2 continued

Phaeophyscia (Ach.) Physcia sciastra, sciastra 17 S, N Moberg P. lithotea Phlyctis boliviensis Nyl. 3 MO

(Fr.) H. Physcia adscendens 7 I, T, E, N Olivier

(Ehrh. ex Physcia aipolia Humb.) 128 I, T, E, N Fürnr.

Physcia americana G. Merr. 48 E

Physcia millegrana Degel. 109 I, N, T

(Tuck.) J. W. Physcia phaea Parmelia phaea 7 Thomson

Physcia stellaris (L.) Nyl. 260 I, S, M

Physcia subtilis Degel. 16

Physcia chloantha, P. luganensis, Physciella chloantha (Ach.) Essl. 76 Phaeophyscia chloantha

Physcia melanchra, Physciella melanchra (Hue) Essl. 23 Phaeophyscia melanchra

Physconia detersa (Nyl.) Poelt Physcia detersa 59 I, T

Physconia enteroxantha (Nyl.) Poelt 3 E, S

Physconia Physconia leucoleiptes (Tuck.) Essl. 65 kurokawae

(Wulfen) Huilia Porpidia albocaerulescens Hertel & albocaerulescens, 9 Knoph Lecidea albocaerulescens,

31 Table 2 continued

L. hebescens

(Scop.) J. Protoblastenia rupestris 9 Steiner

SD, MN, (Orange) H. Macentina Psoraglaena c.f. dictyospora 1 IL, NE, Harada dictyospora MO

(Müll. Arg.) Parmelia bolliana, Punctelia bolliana 157 Krog P. frondifera

Punctelia rudecta (Ach.) Krog Parmelia rudecta 191 N,

(Rehm) R. C. Pyrenula pseudobufonia 25 Harris

Pyxine sorediata (Ach.) Mont. 19

IL, MI, Ramalina Ramalina americana Hale 52 S MN, MO, fastigiata NE, WI

Rinodina efforescens Malme 2 WI, MN

Rinodina freyi H. Magn Rinodina glauca 2

Rinodina populicola H. Magn. 18

Rinodina pyrina (Ach.) Arnold 2

(Stenh.) Bacidia Scoliciosporum chlorococcum 10 I, N Vězda chlorococca

(L.) Otálora, Leptogium Scytinium lichenoides P. M. Jørg. & 41 O lichenoides Wedin

(Nyl.) Arup, Caloplaca Squamulea subsoluta Søchting & 61 subsoluta Frödén

32 Table 2 continued

(Ach.) Conotrema urceolatum 5* Stictis Gilenstam urceolatum

Teloschistes chrysophthalmus (L.) Th. Fr. 105

Thelidium minutulum Körber 1 MO

Coppins & P. WI, MO, Trapelia placodoides 1 James MN, IL (Schrader) Trapeliopsis viridescens Coppins & P. 3 WI James

MO, MN, (L.) F. H. Usnea hirta 2* S-I, O, I SD, NE, Wigg. WI

WI, MN, Verrucaria calkinsiana Servít 4 IL, SD, NE, MO

(Vahl) P. M. Verrucaria furfuracea 2 Jørg.

Verrucaria muralis Ach. 30

Verrucaria nigrescens Pers. 25

Caloplaca Xanthocarpia feracissima H. Magn. 44 feracissima

(Hepp ex Arnold) Xanthomendoza fallax Søchting, Xanthoria fallax 105 S-I, E, S Kärnefelt & S. Kondr.

(Hoffm.) Søchting, Xanthomendoza fulva Xanthoria fulva 51 E Kärnefelt & S. Kondr.

(Räsänen) Søchting, Xanthoria Xanthomendoza ulophyllodes 52 Kärnefelt & ulophyllodes S. Kondr.

Table 3 Macrolichen species that have not been reported since 1960 (data from Iowa Lichen Database - http://www.public.iastate.edu/~herbarium/IowaLichen/index.html) and are likely to be rare in, or extirpated from Iowa. The “#” column represents the total number of accessions of each species. All herbaria that have collections of each species are listed as well as sensitivity ratings if available. : “S” indicates sensitive to sulfur and other air pollutants, “I” indicates species with intermediate sulfur and other air pollutant sensitivity, “T” indicates sulfur and other air pollutant tolerance, “A” indicates acidophytic species, and “O” indicates oligotrophic species. Sensitivity ratings that were contradictory are not included.

Most Recent Sensitivity Genus Species Synonyms # Collection Herbaria Verified

Date Alectoria MIN 23172 nidulifera, WIS Bryoria furcellata 2 1895 WIS, MIN Cornicularia 0040217 fibrillosa arbuscula subsp WIS 11930 Cladonia 1 1894 WIS mitis Cladonia coccifera 2 1895 ISC, NEB ISC 25703 Cladonia NY 33

Cladonia conista 1 1927 NY innominata 01047042 Cladonia cylindrica 2 1957 MSC MSC 52504 NEB, MIN, ISC 25087 Cladonia gracilis 5 1912 T ISC, WIS macilenta var. WIS 37141 Cladonia 1 1902 WIS bacillaris Cladina MIN 19175 Cladonia stellaris 1 1894 MIN stellaris MSC, NEB28575 Cladonia uncialis 5 1957 NEB, MIN, WIS OS, NEB, ISC 344553 Collema pulcellum 3 1895 ISC MICH Collema pustulatum 2 1896 MICH 102566, 102565

Table 3 continued

Enchylium ISC, NEB Synechoblastu MONT, 28840 s ohioense, MSC, 28837 conglomeratum Collema 16 1897 OMA, OS, conglomeratu NEB, MIN, m DUKE Collema MSC 55581 Enchylium polycarpon 2 1957 MSC, OS polycarpon MIN 23258; WIS, MIN, Evernia mesomorpha 4 1895 I WIS 46356; NEB NEB 29390 Anaptychia ISC, MIN, ISC 869929; Heterodermia echinata 4 1895 echinata NEB NEB 29801 Leptogium burnetiae 3 1895 ISC, MIN ISC 25913 ISC, MSC, ISC 321635

Leptogium corticola 20 1903 US, NEB, 34

MIN, WIS MIN 679715 Leptogium saturninum 1 1893 MIN ISC, US, Verified by Sticta Lobaria pulmonaria 7 1901 WIS, NEB, S,O Colbert pulmonaria MIN 2011 MIN 827766 Parmelia squarrosa 2 1913 MIN S,O

Parmotrema michauxianum 1 1897 MIN MIN 895971

MIN 21081 Pertusaria leioplaca 3 1898 MIN,WIS WIS 67065 Pertusaria plittiana 1 1895 ISC ISC 869995 Catapyrenium MIN 655876 tuckermanii, WIS 72014 NEB, Dermatocarpo OMA 13263 Placidium arboreum 8 1898 OMA, n tuckermanii, MIN, WIS Dermatocarpo n arboretum,

Table 3 continued Endocarpon tuckermanii, Endopyrenium tuckermanii MIN 655886

Ramalina sinensis 1 1899 MIN

Collema NEB 31683 plicatile, ISC, NEB, Scytinium plicatile 5 1895 Leptogium MIN plicatile 35 ISC 869935; NY 01399113 Solorina saccata 5 1956 ISC, NY 01399114 0139915 01399116 Cetraria halei, ISC 344492 ISC, US, Tuckermannopsis americana C. ciliaris var. 12 1896* MIN, WIS halei Cetraria US, NEB, MIN 735481 Tuckermannopsis ciliaris 3 1896 A, S ciliaris MIN ISC, WIS, ISC 24902 Usnea angulata 8 1895 NEB, MIN Usnea longissima 1 1894 ISC ISC 321528

Usnea barbata ISC, WIS, ISC 869933 Usnea strigosa 8 1895 A var florida MIN, MIN 652018 Parmelia NEB 38477 Xanthoparmelia conspersa 1 1895 NEB conspersa, P.

Table 3 continued

isidiata

* Erroneously reported in Colbert 2011 as having been last recorded in 1949.

Acknowledgments I thank John Pearson for a great help in gaining permission to collect at all sites. Much appreciation goes to Paul Mayes for taking the time to guide Jim Colbert and I around the Big Sand Mound Preserve. I greatly appreciate all the help from James Lendemer at the New York Botanical Garden for identification of difficult species and all of those that have taught workshops that I have

attended: Peter Nelson, Mark Seaward, David Richardson, Alan Fryday, and James Lendemer. A special thanks goes to the 36 following individuals for helping make collections at Wildcat Den Park, Starr’s Cave State Preserve, and Shimek State Forest: Tracy Chapman, Tom & Deb Podaril, Barb Dotson, Katie Thompson, Catie Steinfadt, Emily Spulak, Joel Bauer, and Samantha Avesing.

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CHAPTER 5 ELEMENTAL CONCENTRATIONS OF IOWA PARMOTREMA SPECIES Published in Evansia 2016 Vol 33 (1) Amy L. Podaril and James T. Colbert Ecology, Evolution, and Organismal Biology Department Iowa State University

Abstract

Eight populations of Parmotrema species from the Shimek State Forest, Iowa (Croton and

Lick Creek Units) were selected for study. Samples of Parmotrema perlatum, P. hypotropum, P. reticulatum, and P. austrosinense were collected and assessed for their concentrations of N, S, P, K, and Fe. We hypothesized that populations near the forest edge might have higher concentrations of these elements. There were no populations of this genus at the forest edge, and no significant differences in elemental concentrations between samples from the middle of the forest or near the forest edge. Three reasons are hypothesized to explain these results: forest age, forest density, and high levels of particulates and gaseous pollutants.

Key Words: Parmotrema, Elements, Deposition, Concentration, Pollution

Introduction

Some lichen species are known to be sensitive to air pollutants as well as nutrient rich conditions (Conti and Cecchetti, 2001). Ammonia, nitrate and phosphate deposition, as well as atmospheric sulfur dioxide concentration are known to cause shifts in lichen communities around the world (Loppi, et al., 1996; Ruoss, 1999; Van Dobben et al, 1996; Van Herk, 2001,

Benner et al., 2007; Benner & Vitousek, 2012; Czeczuga, 1988). Little work has been done on lichen species in response to deposition levels in the Midwest (Bennett et al., 1996,

Bennett & Wetmore, 1997, Bennett & Wetmore, 1999, Bennett & Wetmore, 2003). No studies have focused on Iowa in spite of agricultural practices leading to the highest ammonia emission levels in the Midwest, USA (Stephen & Aneja, 2008). Iowa’s total inorganic nitrogen deposition has increased from 5 kg/ha in 1985 to 8 kg/ha in 2013 (NADP, 2014a).

Atmospheric ammonia concentrations of bordering states lie within levels shown to have significant effects on sensitive species in Europe (NADP, 2012, Van Herk et al., 2003). Since agriculture is the source of 85% of atmospheric ammonia in the United States, it is reasonable to assume that Iowa’s ammonia concentrations are equal or higher than bordering

-3 states (NADP, 2012). Iowa’s NO2 concentrations (18µg m ) may also be of concern as they exceed atmospheric concentrations reported to effect sensitive species in Canada (Gibson et al., 2013), they are however much lower than highly polluted areas such as London (>40µg m-3) which has documented sensitivity of many species (such as Parmotrema perlatum), which is included in this study) (Davies et al., 2007). For some lichen species, the availability of phosphorous can mitigate effects of high nitrogen inputs (Pilkington et al., 2007). As manure and fertilizer use (sources of phosphorus) continues in the state it is important that lichen phosphorus levels are well understood.

There are also some locations in Iowa that have sulfur dioxide concentrations that greatly exceed the National Ambient Air Quality Standard (76 ppb) (IDNR, 2014). Exposure to sulfur dioxide can lead to membrane damage and losses of potassium in lichens (Puckett et al., 1977). According to Lawrey and Hale (1988), lichens with sulfur concentration above

0.20% are rarely observed in unpolluted areas. Wet deposition of SO4 in Iowa in 1992 was much higher than in Minnesota (IA= 14.63 kg/ha, MN=5-8 kg/ha) (NADP, 2014b). This is

43 possibly the reason for a large proportion of the likely extirpated lichen species (85%) in

Iowa which still occur in Minnesota(collected after 1960) (North American Lichen

Consortium Database, 2014). Many areas in Iowa have exceeded the NAAQS for particulate matter in the past (EPA, 2012). Possible sources of particulate matter include agricultural activities, gravel roads, construction worksites, and factories. Iron concentration in lichens has been used to monitor particulates derived from soil/rock and industrial activities (Nash,

2008).

Thirty nine lichens on Iowa’s lichen checklist (Podaril & Colbert, in preparation) are considered sensitive, acidophytic, or oligotrophic species in other studies (Geiser, 2014;

Lawrey, 1993; Richardson, 1992; Wetmore, 1983, 1987, 1989, 1992, 1993a, 1993b, 1995,).

Thirty- three out of Iowa’s 465 species are likely extirpated (haven’t been collected since

1960). Recently, 150 lichen species have been found in the southeast corner of Iowa, 18 of which are new state records (Podaril & Colbert, in preparation). Shimek State Forest contained 100 lichen species and was the richest location out of the five surveyed sites. It also contained populations of various Parmotrema species, which were less common at other sites surveyed. Parmotrema perlatum, is sensitive to SO2 (Smith et al., 2009) and considered oligotrophic in the Pacific Northwest being found at peak frequency at 1.1 kg nitrogen ha-1 yr

-1(McCune & Geiser 2009). This is much lower than Iowa’s nitrogen deposition of 11 kg nitrogen ha-1 yr -1 (NADP, 2014a).

For the current study we measured N, S, P, K, and Fe concentrations in Parmotrema species from Shimek State Forest. We hypothesized that concentrations of these elements would be lower in lichens collected at the center of the forest compared with those growing nearer the edge. The difference was expected to be caused by uptake of elements by the

44 upper canopy of the forest (Krupa, 2003; Fenn et. al, 2013). The collected data reported here from Iowa provide a baseline for future monitoring and increase knowledge about the

Parmotrema habitat in the USA.

Methods

Parmotrema hypotropum, P. perlatum, P. reticulatum, and P. austrosinense were studied at the Croton Unit and Lick Creek Unit of Shimek State Forest. Eight large populations were found and a portion of each was collected for testing. Multiple thalli (minimum of six) were collected on April 26th and 27th, 2014 from each population from an average of three trees. A total of 12 samples, each of a single species were collected from trees at the edge of a trail or creek. One population was collected on a lone tree in the middle of an open grassland area.

Populations that came directly from under an upper canopy or from recently fallen or broken branches are noted in Table 1. No populations of Parmotrema sp. large enough to sample occurred along the forest edge directly bordering agricultural fields.

Once collected, lichen specimens were cleaned of all other plant material (not washed) and dried at 40˚C for 8 hours. Each sample was ground in the Spex CertiPrep

Genogrinder 2000 and then sent to the Soils and Plant Analysis Lab at Iowa State University for analysis of total N (by combustion of material using Leco Truspec CN analyzer and

Elementar Variomax CNS analyzer), S, P, K, and Fe (after microwave digestion are measured on the Spectro Ciros CCD ICP-AES) using methods from Kalra (1998). All samples were split and analyzed in duplicate. One-way ANOVA was performed on

Parmotrema hypotropum collections as this species was found at all sites where the lichens

45 were collected. Populations were grouped to determine if spatial differences in elemental concentrations could be detected.

Results

The average elemental concentrations were 1.42 mg/g for phosphorus, 2.55 mg/g for potassium, 0.69 mg/g for iron, 2.81 mg/g for sulfur and 14.6 mg/g for nitrogen. Iron concentrations in P. reticulatum and P. perlatum from Shimek (0.6 mg/g) are lower than concentrations of these species found in a remote site in southwestern China with relatively clean air (0.8 mg/g) (Zhang et al., 2002) and found in agricultural areas in Italy (1.0- 1.5 mg/g) (Bargali, 1995).

There were no significant differences, for any of the elements tested, between P. hypotropum populations in the interior section of the forest (>200m from closest edge, Figure 1 populations 1,3,6,7) compared to collections made closer to the edge where crop fields are located (Figure 1 populations 2,4,5,8) (p-values >.3). There were also no significant differences between P. hypotropum populations that came from upper canopy branches or from the trunks of trees in open grassland areas (Table 1 Populations 2, 3,7,8) compared to populations from the trunks of trees in the more closed canopy (populations 1, 4, 5, 6) (p- values > .38). There were no significant differences between P. hypotropum populations in the eastern half of the forest (populations 4, 5, 1, 6) compared to the western half of the

Croton Unit (populations 7, 2, 8; p-values > .25) (the single collection from the Lick Creek

Unit was excluded from this statistical analysis).

Figure 1 The overall site is circled in red on the Iowa map. The two units have been magnified with population locations at both the Croton Unit and the Lick Creek Unit are labeled with red points. Numbers correlate with “population” numbers used in Table 1. (Maps produced in ArcGis 10.2)

Table 1 Concentrations of phosphorus (P), potassium (K), iron (Fe), sulfur (S), and percent nitrogen (N) for each population sampled. Upper canopy samples are labeled “UC”. Samples taken from the Lick Creek Unit are labeled “L”. Open grassland samples are labeled “OG”. All concentrations are an average of the duplicate measurements from each site. Site numbers correspond to numbers used in Figure 1.

Population Species P K Fe S N

(mg/g) (mg/g) (mg/g) (mg/g) (mg/g)

1 P. hypotropum 0.8 2.1 0.8 2.7 13.9

2UC P. hypotropum 1.7 2.9 0.7 2.8 17.0

P. perlatum 1.4 2.5 0.8 2.9 14.9

3-L/UC P. hypotropum 1.5 2.3 0.5 2.5 13.8

4 P. hypotropum 1.0 2.3 0.7 2.5 14.0

P. hypotropum 1.9 3.0 0.6 2.9 13.7

P. perlatum 1.2 2.2 0.7 2.9 12.6

6 P. reticulatum 1.6 2.7 0.6 3.2 16.3

P. hypotropum 1.3 2.5 0.5 3.1 14.7

7UC P. hypotropum 1.6 3.2 0.7 2.5 13.7

8 OG P. hypotropum 1.1 2.1 0.9 3.3 14.8

P. austrosinense 2.0 2.7 0.8 2.7 16.2

AVERAGE 1.4 2.5 0.7 2.8 14.6

Discussion

It is difficult to draw conclusions from such a small sample size and lack of other studies as to whether these elemental levels are high or low values for Parmotrema species.

Only two studies were found to report elemental concentrations in the genus. The sulfur concentration of transplanted P. austrosinense in Argentina was ~1.75 mg/g after six months in an urban site and ~1.0 mg/g after six months in an unpolluted site (Cañas et al., 1997,

Zhang et al., 2002). These values are low compared to the sulphur concention of P. austrosinense found at Shimek State Forest (2.7 mg/g).

The lack of significant differences found in elemental concentrations from the center and near the edge of the forest could be the result of at least two factors. First, it is possible that the gaseous and particulate inputs are higher than could be absorbed by the upper canopy. Second, the forest may not be dense enough to have a filtering effect on the levels of

N, S, P, Fe, and K found in the lichens. This could be related to the age of the forest or the presence of wide man-made trails that increase air circulation through the trees. Indeed, the more open areas in this forest unit seemed to be favored by the Parmotrema species. It is also important to note that there were no large populations of Parmotrema found close to the edges of the forest which directly faced crop fields. It is possible that the populations thrived only in areas that were partially filtered by the forest canopy.

Iowa includes many eutrophic lichen habitats due to nutrient inputs from agricultural practices, vehicle exhaust, and other anthropogenic activities. The studied forest has a large number of rare, or potentially extirpated, lichen species, and it is crucial that the current populations are managed and monitored to prevent further losses. Attempts to reintroduce

(e.g., Sillet et al., 2000) extirpated lichen species (e.g., Lobaria pulmonaria, Colbert, 2011)

49 could be unsuccessful due to the current deposition levels of particulates and atmospheric pollutants.

Acknowledgments

I thank the Lois H. Tiffany Scholarship Committee for choosing to fund our work. I also thank Toby Spribille for his advice on lichen grinding methods and David Richardson for providing a very helpful review.

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CHAPTER 6 SUMMARY AND CONCLUSIONS

Although study of Iowa lichens began in the late 1800’s, there is still much left to learn about the diversity and distribution of lichens of Iowa. For this study nearly 400 lichen collections were made in southeast Iowa. These collections have begun to expand our understanding of lichen diversity in a part of the state that had previously been poorly explored (Figure1). A total of 150 lichen species were identified, 18 of which were new state records. The majority (78%) of these new records were crustose lichens. Due to the lack of historical records for crustose species, it is unknown whether the new state record crustose lichen species have recently migrated into the area or merely were overlooked in the past.

These 18 species have been added to the Iowa Lichen Checklist

(http://www.public.iastate.edu/~herbarium/IowaLichen/index.html), which now totals 465 species for the state. Based on lichen diversity estimates from surrounding states this appears to be a reasonable estimate for the State of Iowa.

Figure 3 Map of all Iowa lichen collections before our study (left) and all Iowa lichen collections made including our study (right). Each point represents a single collection. Since old collections didn’t have georeferenced data, all collections were randomly placed within the county it was collected in. Therefore the exact position of each point does not correspond to the precise collection location. The year the collection was made is indicated by the corresponding color in the legend.

Although we now have a reasonable estimate of the lichen diversity in the State of Iowa, we continue to have a very limited understanding of the spatial distribution of lichen species within the State of Iowa. Some species, e.g., Physcia stellaris, clearly seem to have a state- wide distribution. We have a much poorer understanding of whether some Iowa lichen species show more restricted distribution patterns within the state. For example, Teloschistes chrysophthalmus, which is fairly common in some parts of the state, appeared to be quite rare in the four southeast Iowa counties (3 accessions) studied here. Is T. chrysophthalmus found throughout the State of Iowa? Is this species more common in certain regions of the state?

Systematic attempts to collect particular species in various regions of the state would be required to answer such questions.

Of the 465 species currently on the Iowa Lichen Checklist, 33 species (~ 7%) are considered here to be potentially extirpated. The current study found one species (Cladonia scabriuscula) that was previously listed as potentially extirpated (Colbert, 2011), however since only a single collection was made, this species should still be considered to be rare within the state. This represents a significant loss of biological diversity over the approximately 130 years of sporadic attention to Iowa’s lichen diversity. This may well be an underestimate since insufficient data are available to make any estimate of the number of potentially extirpated microlichens in the state. Although this is a substantial loss of biodiversity, it is smaller than the loss of diversity observed in other groups of organisms in

Iowa over approximately the same time frame, e.g., the 50% loss of species diversity in freshwater mussels since 1900 (Poole and Downing, 2004). At this stage it is impossible to predict whether Iowa’s lichen diversity will continue to decline, or whether, as a result of habitat or climate change, additional species will be able to establish themselves in the state.

Given that nine of Iowa’s 99 counties have never had any lichen collections made, and the relatively small total number of accessions of Iowa lichens (~ 8,000) that we are currently aware of, it is also possible that additional collecting within the state will lead to an increase in the number of lichen species known to be present in Iowa.

There could be numerous causes of the potential extirpation of 33 species of lichens from Iowa, but the two most likely factors are habitat loss and high nitrogen deposition.

From our elemental analysis study there were no significant differences in N, P, S, K, and Fe concentrations found in Parmotrema species from various locations within Shimek State

Forest. Any attempts to reintroduce (e.g., Sillet et al., 2000) extirpated lichen species (e.g.,

Lobaria pulmonaria, Colbert, 2011) will need to consider to the possibility of atmospheric deposition of pollutants.

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APPENDIX

IOWA LICHEN CHECKLIST 2015

Iowa Lichen Checklist, 2015. New state record species have * in the “Found in SE IA” column. Online version available at Iowa Lichen Project http://www.public.iastate.edu/~herbarium/IowaLichen/index.html

Most Found # of Recent Sensitivit Surroundin Genus Species Synonyms in SE Accession Herbaria Comments Collectio y g States IA s n Date

Acarospora KANU, Acarospora americana 4 2007 cinereoalba MICH, NY Acarospora cervina 4 1894 F, MICH

Acarospora citrina 3 1978 ISU KANU, Acarospora contigua 4 2006 UCR Acarospora oligospora 1 1903 MICH KANU, Acarospora veronensis 13 2003 MIN, FH, F, MICH MICH, Acrocordia cavata Arthopyrenia cavata Y 8 2013 MIN, ISC, NY, WIS Arthopyrenia gemmata, Acrocordia gemmata 3 1972 ISC, MIN A. alba, A. sphaeroides

Arthopyrenia KANU, Acrocordia megalospora macrospora, A. finkii, 9 2009 MIN,MSC Pyrenula megalospora

ISC, NEB, WIS, TLE, BRY ASU, Opegrapha diaphora, Alyxoria varia Y 87 2013 MSC, O. varia TENN, NY, KANU, MIN, Rinodina dakotensis, R. KANU, finkii, R. inaequalis, R. MIN, BRY, Amandinea dakotensis pennsylvanica, R. Y 30 2013 DRAKE, subplumbea, R. ISC, WIS, subpyriniformis MSC,

Amandinea milliaria Rinodina milliaria 8 1969 ISC Buellia polyspora, KANU, Amandinea polyspora Buellia punctata var. 7 2002 MIN polyspora

ISC, 63 Buellia punctata, B. KANU,MIN Amandinea punctata Y 65 2013 n myriocarpa, B. pullata , MSC, WIS, NEB

MIN, ISC, Anaptychia palmulata A. "palmatula" auct. 8 1978 MICH, us Arthopyrenia biformis, A. parvula, A. conformis auct. N. Am., Anisomeridium biforme Y 4 2002 ISC, KANU Ditremis biformis, Trimmatothele umbellulariae Anisomeridium ISC, KANU, nyssaegenum, Ditremis Anisomeridium polypori Y 12 2013 MSC, MIN, nyssigena, NY Arthopyrenia willeyana

Arthonia atra Opegrapha atra Y 2 2013 ISC I

NY, MSC, KANU, ISC, Arthonia dispersa 33 2003 MIN, MICH, US, NEB (+)Arthonia glaucella 1 1965 SBBG KANU, US, Arthonia patellulata Y 9 2013 ISC

MIN, ISC, Arthonia polymorpha 6 1895 NEB

MSC, ISC, ASU, MIN, DUKE, Arthonia punctiformis 38 2001 NEB, WIS

64 ,BRY, SBBG, OMA ISC, KANU, MSC, NY, Arthonia radiata Y 49 2013 SRP, NEB, I, a MIN, WIS, DUKE

ISC, NEB, Arthopyrenia cerasi Pyrenula cerasi 5 1894 DUKE KANU, Arthothelium ruanum 2 2003 MIN ASU, DRAKE, ISC, MSC, OMA, Arthothelium spectabile Arthonia spectabilis Y 34 2013 SBBG, NEB, MIN, DUKE, WIS, BRY

KANU, Bacidia populorum, B. MICH, NY, Arthrosporum populorum acclinis, Bilimbia 15 2003 NEB, acclinis MIN,WIS

ISC, KANU, Aspicilia cinerea Lecanora cinerea 13 2002 NEB, MIN, NY, WIS

Aspicilia limitata Lecanora limitata 1 2000 KANU ISC, KANU, MICH, Athallia holocarpa Caloplaca holocarpa Y 44 2013 n NEB, MIN, BRY MSC, Athallia pyracea Caloplaca pyracea 3 2006 KANU, WIS

MIN, 65 Athallia vitellinula Caloplaca vitellinula 3 2003 I-T KANU

ISC, KANU, MICH, Bacidia bagliettoana B. muscorum 25 2011 NEB, MIN, WIS KANU, Bacidia circumspecta 7 2009 OMA, NY ISC, KANU, MICH, Bilimbia pammellii, MONT, Bacidia coprodes Bilimbia granosa, Y 47 2013 MSC, OSC, Bilimbia trachona auct. MIN, NY WIS, NEB, DUKE Bacidia laurocerasi 1 1894 ISC

ISC, MICH, MSC, OMA, Bacidia polychroa 26 1903 NEB, MIN, WIS

ASU, ISC, KANU, Bacidia rubella Y 12 2013 MICH, I MSC, NEB, MN,

Bacidia schweinitzii Biatora leucampyx Y 3 2012 MIN, ISC

ISC, MICH, MSC, OSC, Bacidia suffusa Y 26 2013 NEB, MIN, DUKE

Bacidia assulata, B. Bacidina assulata 1 1894 MICH effusa auct. Bacidina egenula Bacidia egenula Y 10 2013 ISC, KANU,

ISC, MICH, OSC, NEB, Bacidina inundata Bacidia inundata Y 41 2013

MIN, BRY, 66 DUKE

Bagliettoa calciseda Verrucaria calciseda Y 3 2013 ISC NEB, MIN, Biatorella hemisphaerica Biatora fossarum 4 1895 WIS,

Bacidia sabuletorum, ISC, KANU, B. hypnophila, MICH, Mycobilimbia Bilimbia sabuletorum Y 40 2013 MSC, WIS, sabuletorum, NEB, MIN, Myxobilimbia UPS, BRY sabuletorum

Blastenia ferruginea Caloplaca ferruninea 5 1896 NEB Collema cheilum, C. Blennothallia crispa 4 1978 ISC crispum Alectoria Bryoria chalybeiformis 1 1893 NEB chalybeiformis Alectoria nidulifera, Bryoria furcellata 3 1895 WIS, MIN Cornicularia fibrillosa

Amandinea turgescens, KANU, (#)Buellia badia Buellia turgescens, 7 2008 MSC, NEB, Buellia turgescentoides MIN, ISC Hafellia disciformis, B. ISC, NEB, Buellia disciformis 14 1963 parasema MIN, MIN, WIS, Buellia erubescens Buellia stillingiana 7 1957 MSC Buellia maculata Buellia stigmaea 2 1996 MIN, ASU Buellia sequax 1 2007 KANU ISC, NEB, Buellia spuria 9 1903 MIN, WIS, Calicium lenticulare 3 1903 ISC Calicium parietinum 3 NA NEB

NEB, MSC, 67 Calogaya lobulata Caloplaca lobulata 3 1965

DUKE

Calogaya pusilla Caloplaca pusilla 2 1903 MIN

Caloplaca ahtii 6 1992 MIN, ISC Caloplaca atroalba Lecania perproxima 3 2009 ISC Caloplaca camptidia 2 1964 ISC ASU, DUKE, ISC, KANU, Caloplaca cerina Y 200 2013 MICH, S-I MSC, UPS, WIS, NEB, MIN, ISC, DUKE, Caloplaca chlorina 4 2012 KANU Placodium ISC, DUKE, Caloplaca cinnabarina 14 1970 cinnabarinum NEB (*)Caloplaca grimmiae 1 2002 KANU

DUKE, ISC, KANU, MOR, Caloplaca microphyllina Y 73 2013 OMA, NEB, MIN, WIS, BRY, MIN, MSC, Caloplaca pollinii 10 1961 WIS MIN, Caloplaca pratensis 4 2000 KANU Caloplaca saxicola 3 1992 MIN, ASU ISC, KANU, Caloplaca soralifera Y 6 2012 MIN ISC, KANU, Caloplaca ulcerosa Y 11 2013 MIN

WIS, UPS, Caloplaca ulmorum 4 2006 KANU ASU, ISC, KANU, MOR, MSC, S-I, tt,s- Candelaria concolor Y 200 2013 OMA, NEB, I,e,n MIN, WIS, BRY, ISC, KANU, Candelaria fibrosa Y 66 2013 MSC, OMA, MIN, WIS,

Candelariella aurella Y 7 2013 ISC, MIN n ASU, MIN, Candelariella efflorescens Y 9 1996 NY, KANU, ISC KANU, Candelariella subdeflexa 7 2003 MIN KANU, ISC, Candelariella vitellina 29 2007 SBBG, I, tt, n NEB, MIN

KANU, Candelariella xanthostigma 8 2009 MOR, MSC, S-I, n MIN Dermatocarpon Catapyrenium cinereum cinereum, D. 19 1977 ISC, NEB hepaticum Catillaria chalybeia Y 2 2013 ISC

With Teloschistes Catillaria glauconigrans Y* 1 2013 ISC WI, SD chrysophthalmu s #873384 Bacidia declinis, Catillaria nigroclavata Bilimbia declinis, Y 4 2010 ISC, KANU Lecidea declinis Chaenotheca furfuracea Coniocybe furfuracea 3 2011 ISC

MICH, ISC, Allarthonia caesia, KANU, 69 Chrysothrix caesia Arthonia lecideella, Y 93 2013 MSC, US, Arthonia caesia NEB, MIN, NY, BRY

Chrysothrix xanthina 1 2011 ISC Aspicilia caesiocinerea, Circinaria caesiocinerea 2 2000 KANU, WIS Lecanora caesiocinerea Aspicilla calcarea ; ISC, MSC, Circinaria calcarea 11 1895 Lecanora calcarea OMA, NEB Aspicillia contorta ; MIN, WIS, Circinaria contorta 16 1992 Lecanora contorta NY, DUKE Aspicillia gibbosa ; Circinaria gibbosa Lecanora gibbosa, L. 4 1894 MIN, WIS gibbosula

Cladonia apodocarpa Y 9 2013 ISC

ISC, MIN, Cladonia arbuscula Cladina arbuscula 4 1964 ss,ii NEB Cladonia arbuscula subsp mitis Cladina mitis 1 1894 WIS Cladonia botrytes 2 1961 MIN, ISC Cladonia caespiticia Y 5 2012 ISC, MIN ISC, KANU, Cladonia cariosa Y 25 2013 MOR, MIN ISC, NEB, Cladonia cervicornis 17 1993 MIN

BDI, ISC, MSC, NY, Cladonia chlorophaea Y 43 2013 ss,ii,e NEB, MIN, WIS, BRY,

70 Cladonia coccifera 2 1895 ISC, NEB

ISC, NEB, Cladonia coniocraea Y 26 2013 I,ss MIN, WIS Cladonia conista Cladonia innominata 2 1923 NY ISC, NEB, Cladonia cristatella Y 32 2013 I MIN, WIS ISC, MSC, Cladonia cryptochlorophaea Y 6 2013 NEB, NEB Cladonia cylindrica 2 1957 MSC ISC, KANU, Cladonia decorticata Y 7 2013 OMA ISC, NEB, Cladonia didyma 10 2007 MIN, WIS ISC, NEB, Cladonia fimbriata Y 14 2013 S-I, ss,e MIN, WIS

ISC, KANU, OMA, NEB, Cladonia furcata Y 73 2013 ii, o, tt MIN, WIS, BRY

NEB, MIN, Cladonia gracilis 5 1912 tt ISC, WIS Cladonia humilis 2 1961 MIN ISC, KANU, Cladonia macilenta Y 23 2013 MSC, NEB, ss, e, P MIN macilenta var. Cladonia 1 1902 WIS bacillaris Cladonia magyarica 1 2007 ISC Cladonia merochlorophaea 1 1986 OMA

Cladonia multiformis 2 1978 ISC, NEB COE, ISC, MSC, OMA, 71 Cladonia parasitica Y 35 2013 TENN, NEB, MIN, WIS ISC, MSC, OMA, NEB, Cladonia peziziformis C. mitrula, C. capitata Y 55 2013 MIN, WIS, BRY

ISC, MIN, Cladonia pleurota 8 2008 MSC COE, ISC, Cladonia pyxidata Y 44 2013 KANU, m NEB, MIN

ASU, COE, ISC, MSC, Cladonia rangiferina Cladina rangiferina 24 2013 ss,ii NEB, MIN, WIS

KANU, Cladonia rei Y 17 2013 MSC, MIN, ISC, KANU, Cladonia robbinsii Y 10 2013 NEB, MIN Was previously Cladonia scabriuscula Y 3 2013 ISC, MIN on ext. list

COE, ISC, KANU, Cladonia squamosa Y 21 2012 ii,e MSC, MIN, WIS, BRY

Cladonia stellaris Cladina stellaris 1 1894 MIN WI, IL, MO Cladonia strepsilis Y* 1 2012 ISC C. clavulifera, C. WI, IL, NE, Cladonia subcariosa polycarpia, C. Y 8 2013 ISC MO polycarpoides

Cladonia subtenuis Y 12 2013 ISC, KANU Cladonia subulata Y* 2 2012 ISC MN, IL

Cladonia symphycarpa Y 19 2013 ISC, MIN

MSC, NEB, Cladonia uncialis 5 1957 MIN, WIS Cladonia verticillata 1 NA OS

Clavascidium umbrinum Placidium umbrinum 1 2001 KANU Dimerella pineti, D. WI, SD, MO, Coenogonium pineti diluta, Microphiale Y* 1 2013 ISC MN, IL diluta Synechoblastus US, NEB, Was previously Collema nigrescens 5 2008 nigrescens MIN, ISC on ext. list ISC,OS, Collema pulcellum 3 1895 NEB Collema pustulatum 1 1896 MICH

ISC, MSC, Collema subflaccidum Y 13 2013 S MIN, NEB COE, ISC, KANU, MONT, Collema tenax Collema pulposum 41 2009 MSC, OS, NEB, MIN, DUKE, BRY Synechoblastus Collema texanum Y 1 NA ISC texanus, S. laciniatus ISC, NEB, Cyphelium tigillare Y 26 2012 OS, MIN (*)Dactylospora lurida 1 1991 NY

Leptogidium intricatulum,

Dendriscocaulon intricatulum 1 2002 KANU 73 Polychidium intricatulum

Dermatocarpon dolomiticum 2 1987 NY

Dermatocarpon luridum Y 5 2013 ISC, MIN

DUKE, ISC, KANU, MSC, OMA, Dermatocarpon miniatum Y 107 2013 OS, WIS, NEB, MIN, BRY

Dermatocarpon muehlenbergii 1 1961 DUKE

Rinodina oreina, R. ISC, KANU, hueana, R. Dimelaena oreina 35 2003 MSC, NEB, novomexicana, R. MIN, WIS suboreina

Urceolaria NEB, MIN, Diploschistes actinostomus 3 1899 actinostoma WIS

ISC, KANU, Diploschistes muscorum Y 12 2013 MIN, NY

ASU, ISC, MSC, OMA, Diploschistes scruposus Urceolaria scruposa 32 1982 OS, WIS, NEB, MIN, NY ISC, MONT, MSC, OMA, Buellia alboatra, Diplotomma alboatrum 21 2008 OS, NEB, Rhizocarpon alboatrum MIN, WISE, BRY

Distopyrenis americana 4 1895 NY Collema bachmanianum, ISC, MICH, Enchylium bachmanianum 17 1983 74 Collemodes MIN, COE bachmanianum Collema ISC, MONT, conglomeratum, MSC, OMA, Enchylium conglomeratum Synechoblastus 16 1897 OS, NEB, ohioense, S. cyrtaspis, MIN, DUKE S. pycnocarpus Collema limosum, C. Enchylium limosum 3 1961 MIN, WIS glauconigrescens Collema polycarpon, Synechoblastus Enchylium polycarpon 2 1957 MSC, OS polycarpus, S. wyomingensis Staurothele MIN, ISC, Endocarpon diffractellum diffractella, Verrucaria Y 7 2013 WIS diffractella Endocarpon pallidulum 12 2011 KANU, NY Endocarpon petrolepideum 1 2003 KANU

ASU, ISC, MSC, OMA, Endocarpon pusillum Y 78 2013 WIS, NEB, MIN, BRY

Endocarpon subnitescens 1 1989 SBBG Pyrenula leucoplaca Eopyrenula intermedia Y 17 2013 MIN, NY var. pluriloculata Euopsis pulvinata Pyrenopsis pulvinata 2 1894 ISC, NEB Evernia mesomorpha 5 1895 WIS, MIN I, ii Parmelia baltimorensis, ISC, KANU, Flavoparmelia baltimorensis Y 5 2013 Pseudoparmelia MIN baltimorensis Parmelia caperata, P. ISC, BDI, cylisphora, P. KANU, 75 flavicans, P. herreana, MSC, OMA, Flavoparmelia caperata Y 123 2013 I, e, n P. negativa, US, WIS, Pseudoparmelia NEB, MIN, caperata BRY ISC, MSC, Flavoplaca citrina Caloplaca citrina Y 29 2013 n MIN, WIS Parmelia flaventior, P. andreana, P. ISU, MIN, Flavopunctelia flaventior 10 2006 e, n kernstockii, Punctelia KANU flaventior Parmelia soredica, P. ulophyllodes, P. Flavopunctelia soredica 3 1978 ISC manshurica, Punctelia soredica ISC, ASU, KANU, MSC, OMA, OS, TENN, Graphis scripta Y 159 2013 I WIS, NEB, MIN, SBBG, NY, BRY

ASU, ISC, KANU, Caloplaca aurantiaca, Gyalolechia flavorubescens Y 107 2012 MSC, OMA, C. flavorubescens NEB, MIN, WIS DUKE, ASU, ISC, Caloplaca erythrella, KANU, Gyalolechia flavovirescens Y 71 2013 C. flavovirescens MSC, NEB, MIN, WIS, BRY Caloplaca bryochrysion, C. Gyalolechia xanthostigmoidea 3 1963 ISC epiphyta, C. xanthostigmoidea Heppia adglutinata 4 1894 NY

WIS, ASU,

ISC, KANU, 7

Heppia lutosa 36 2007 MSC, OMA, 6 OS, UPS, NEB, MIN,

ISC, MIN, Heterodermia echinata Anaptychia echinata 4 1895 NEB

ISC, NEB, Heterodermia hypoleuca Anaptychia hypoleuca 7 1978 MIN

Anaptychia heterochroa, A. Heterodermia obscurata hypoleuca var. 1 1978 ISC colorata, A. obscurata, A. sorediifera Anaptychia ISC, MSC, Heterodermia speciosa pseudospeciosa var. 32 2007 NEB, MIN, tremulans, A. speciosa WIS, Ionaspis epulotica, Hymenelia epulotica 2 1961 ISC Lecanora epulotica

ISC, KANU, Physcia adglutinata, P. MOR, MSC, Hyperphyscia adglutinata elaeina, Physciopsis Y 53 2011 I, a NEB, MIN, adglutinata, P. elaeina NY KANU, Hyperphyscia confusa 11 2011 TLE, ISC, KANU, Physcia syncolla, MSC, NEB, Hyperphyscia syncolla Y 134 2013 Physciopsis syncolla MIN, WIS, BRY

Ionaspis alba 1 2008 ISC Polyblastiopsis ISC, MSC, (+)Julella fallaciosa 13 2012 fallaciosa WIS, NY Collema furvum, C. Lathagrium fuscovirens fuscovirens, C. 3 1992 MIN, WIS

tuniforme 77

Lecania croatica Y* 1 2013 ISC Lecania cyrtella Y* 1 2013 ISC a SD, MN ISC, NEB, Lecania erysibe Y 6 2013 MIN Bacidia naegelii, Lecania naegelii 6 1968 ISC, MICH Bilimbia naegelii Lecania nylanderiana 1 1894 NEB Bacidia sibiriensis, B. Lecania subfuscula 1 1989 SBBG subfuscula Lecanidion atratum 1 2007 KANU ISC, NEB, Lecanora albella L. pallida 7 1895 MIN, WIS, ISC, SBBG, Lecanora albellula L. piniperda 4 2011 MIN ISC, NEB, Lecanora allophana L. subfusca 16 2010 I MIN, WIS,

Lecanora appalachensis Y* 1 2013 ISC KANU, Lecanora argentata 15 2000 NEB, MIN, Lecanora argopholis 1 1897 MIN caesiorubella subsp. Lecanora 2 1895 WIS caesiorubella Lecanora cenisia 1 2007 ISC ISC, MSC, Lecanora chlarotera L. rugosella Y 9 2013 WIS, MIN, I BRY

Lecanora circumborealis 2 1895 MIN, WIS I DUKE, ISC, KANU,

78 Lecanora dispersa Y 40 2013 MSC, WIS, T, n NEB, MIN, BRY Lecanora fuscescens Lecidea fuscenscens 1 1961 ISC ISC, KANU, Lecanora hagenii 12 2011 T NEB, MIN ISC, KANU Lecanora hybocarpa 7 2011 MIN, Lecanora hypoptoides 1 1960 ISC

Lecanora invadens 1 1961 MIN Lecanora iowensis 2 1963 ISC, MICH ISC, KANU , MONT, Protoparmeliopsis Lecanora muralis 35 2003 MSC, NY, T, n muralis NEB, MIN, WIS, muralis var. Lecanora 1 1894 WIS versicolor Lecanora opiniconensis 4 1897 CANL, MIN

Lecanora perpruinosa 2 1894 MIN Lecanora pulicaris 1 1895 NEB a ISC, MSC, Lecanora saligna Lecanoropsis saligna 11 2013 WIS, NEB, I MIN, BRY MSC, MIN, Lecanora semipallida Lecanora flotoviana 8 1903 WIS ISC, KANU Lecanora strobilina Y 17 2013 MIN, ISC, MIN, Lecanora symmicta Lecidea symmicta Y 21 2013 WIS, MOR ISC, KANU, Lecanora thysanophora Y 15 2014 MIN, KANU, Lecanora valesiaca 3 2003 MIN

79 ISC, NEB, Lecanora varia 8 2001 MIN, WIS, ISC, NEB, Lecidea carnulenta 5 1903 MIN, WIS ISC, NEB, Lecidea plebeja L. myriocarpoides 9 1911 MIN, WIS, ISC, NEB, Lecidea varians Pyrrhospora varians 8 1894 WIS, MIN, KANU

Lecidea elaeochroma, Lecidella elaeochroma L. olivacea, L. limitata 2 NA WIS auct. Lecidea euphorea, L. Lecidella euphorea 9 1916 MIN, WIS glomerulosa

Bacidia arthoniza, Lecidea micacea, L. Lecidella stigmatea 1 1963 MIN stigmatea, L. vulgata, L. lacus-crateris

Lecidea chalybeiformis, L. Leimonis erratica erratica, L. 3 2012 ISC suberratica, Micarea erratica Lepraria caesiella 4 2013 ISC DUKE, ISC, KANU, Crocynia aliciae, C. MICH, Lepraria finkii americana, L. Y 82 2013 a MOR, US, lobificans MIN, WIS, MSC Leproloma membranaceum, Lepraria membranacea 25 1986 ISC, OMA Amphiloma lanuginosum Lepraria normandinioides 6 2012 ISC Leproplaca cirrochroa Caloplaca cirrochroa 6 1992 MIN 1 2008 ISC Leptogium azureum 80

Brodo lists as Leptogium burnetiae 3 1895 ISC, MIN "uncommon" DUKE, ISC, MONT, Leptogium chloromelum 10 1961 NEB, MIN, WIS

ISC, MSC, Leptogium corticola 20 1903 US, NEB, MIN, WIS

ISC, US, Leptogium cyanescens 19 2011 ss, sss MIN, WIS Leptogium hirsutum 3 NA WIS Leptogium saturninum 1 1893 MIN Leptorhaphis atomaria 3 1961 ASU, MSC (#)Leptorhaphis epidermidis 1 1962 ISC

Gonohymenia cribellifera, Lichinella cribellifera 3 1963 MIN Rechingeria cribellifera Gonohymenia Lichinella nigritella nigritella, Thyrea 1 NA MIN nigritella Arthopyrenia hyalospora, ISC, MIN, Lithothelium hyalosporum Plagiocarpa 3 1895 NEB hyalospora, Pleurotrema solivagum ISC, US, Lobaria pulmonaria Sticta pulmonaria 7 1901 WIS, NEB, ss, O, S MIN Loxospora elatina Haematomma elatinum 1 NA WIS Haematomma Loxospora ochrophaea 4 1897 MIN, NY ochrophaeum 81

Melanelia subaurifera, ISC, WIS, Melanelixia subaurifera 8 1978 e, s-i Parmelia subaurifera MIN Melanelia exasperata, US, NEB, Melanohalea exasperata Parmelia exasperata, 8 1896 tt,ii,m MIN, WIS P. aspera, P. aspidota

Melaspilea arthonioides 1 1894 WIS ISC, MIN, Melaspilea demissa 6 1968 WIS Bacidia trisepta, Micarea peliocarpa 3 1982 ISC Bilimbia trisepta Micarea prasina Catillaria prasina 3 2003 KANU (*)Muelleralla lichenicola 1 1991 NY Sphinctrinella (+)Mycocalicium calicioides 3 1896 MIN, NY calicioides ISC, MONT, M. parietinum, (+)Mycocalicium subtile 8 2012 MSC, MIN, Calicium subtile NY

CANL, (+)Mycoglaena quercicola 9 1895 MSC, US, MIN, WIS, (+)Mycomicrotheli wallrothii 3 1895 WIS a Dermatina Mycoporum compositum 3 1962 ISC, MIN pyrenocarpa Parmelina aurulenta, ISC, KANU, Myelochroa aurulenta Parmelia aurulenta, P. Y 77 2013 MSC, NEB, silvestris MIN, BRY

Parmelina galbina, Parmelia galbina, P. ISC, MIN, Myelochroa galbina Y 37 2013 subquercifolia, P. WIS, NEB sulphurosa Acarospora amphibola, A. smaragdula, A. Myriospora smaragdula smaragdula var. 5 1962 ISC, OMA lesdainii, Silobia smaragdula 82 Verrucaria punctiformis; NEB, MSC, (#)Naetrocymbe punctiformis 11 1963 Arthopyrenia MIN, ISC punctiformis (*)Opegrapha anomea Opegrapha quaternella 1 1894 NEB Opegrapha herbarum Y 4 2013 ISC OMA, ISC, Opegraphoidea (*)Opegrapha pulvinata 16 1898 MIN, NEB, pulvinata NY, WIS

Opegrapha vulgata Y 2 1968 ISC I ISC, US, Parmelia saxatilis 10 1980 I, tt, e, ii, a NEB, WIS Parmelia squarrosa 2 1913 MIN S,ss,o sss ISC, KANU, I-T, MT, Parmelia sulcata 27 2000 MSC, US, e, i-t, ttt MIN

Parmotrema arnoldii Parmelia arnoldii 1 2012 ISC ii, o Parmelia Parmotrema austrosinense Y* 1 2013 ISC WI, MO, IL austrosinensis ISC, MSC, Parmelia crinita, Parmotrema crinitum Y 20 2013 US, NEB, ss,o Parmelia proboscidea MIN, WIS Parmelia hypotropa, Parmelia hypotropa ISC, MIN, Parmotrema hypotropum var. sorediata , P. Y 13 2013 ttt NEB, WIS cetrata var. hypotropoides ISC, KANU, Parmotrema margaritatum Parmelia margaritata 36 2011 MSC, US, NEB, MIN,

Parmotrema michauxianum 1 1897 MIN 895971 Parmelia perforata, P. ISC, NEB, Parmotrema perforatum erecta, P. 7 1978 ss, o, ii 83 MIN

hypotropoides

Parmelia perlata, P. Parmotrema perlatum trichotera, Y 5 2013 ISC, WIS a Parmotrema chinense

Parmelia reticulata, ISC, MIN, Parmotrema reticulatum Y 17 2013 S Rimelia reticulata KANU Parmelia michauxiana, Parmotrema submarginale 1 1978 ISC P. epiclada

Canomaculina subtinctoria, Rimeliella ISC, MSC, Parmotrema subtinctorium 11 1978 subtinctoria, Parmelia US, MIN subtinctoria

ASU, ISC, Peltigera canina Y 82 1984 NEB, MIN, tt WIS, BRY ISC, NEB, Peltigera didactyla 21 1979 ss MIN, WIS

ISC, MIN, Peltigera elisabethae 8 1961 CINC, NEB ISC, MSC, Peltigera evansiana 26 2011 NEB, MIN, WIS ISC, MSC, Was previously Peltigera horizontalis 5 2012 I, ii WIS, MIN, on ext. list Peltigera lepidophora 2 1962 MIN Peltigera neckeri 6 1962 MIN

TENN, ASU, ISC, Peltigera polydactylon 31 2012 MSC, NEB, MIN, WIS

ISC, KANU, Peltigera praetextata Y 37 2013 NEB, MIN, ss

WIS, HARV 84

ISC, KANU, Peltigera rufescens 23 2011 MSC, NEB, MIN, WIS

Pertusaria consocians 2 1993 SBBG Pertusaria leioplaca 2 1898 MIN, WIS With Physcia WI, MN, Pertusaria macounii Melanaria macounii Y* 1 2012 ISC americana # MO 872089

Pertusaria plittiana 1 1895 ISC ISC, KANU, Pertusaria pustulata Y 41 2013 MSC, NEB, MIN, WIS

Pertusaria rubefacta Y 1 2013 ISC (+)Phaeocalicium curtisii Calicium curtisii 3 1996 ASU, MIN Phaeographis inusta 1 1963 ISC

ISC, KANU, Phaeophyscia adiastola Physcia adiastola Y 57 2013 TLE, MIN, ISC, COE, KANU, Physcia ciliata, P. Phaeophyscia ciliata Y 139 2013 MSC, WIS, obscura auct. NEB, MIN, BRY Phaeophyscia ISC, KANU, Phaeophyscia hirsuta cernohorskyi, Physcia Y 57 2013 MIN, hirsuta

ISC, OMA, Phaeophyscia hirtella Y 20 2012 MIN, KANU, WIS Physcia hispidula, P. Phaeophyscia hispidula Y 19 2013 ISC, MIN setosa Phaeophyscia insignis 7 2009 KANU

ISC, MSC,

OMA, WIS, 85 Physcia orbicularis, P.

Phaeophyscia orbicularis Y 40 1979 NEB, MIN, I, ii,e,n virella KANU, BRY

ISC, KANU, Phaeophyscia pusilloides Physcia pusilloides Y 29 2013 SBBG, TLE, MIN

Physcia orbicularis f. ISC, KANU, rubropulchra, P. Phaeophyscia rubropulchra Y 64 2012 MSC, NEB, n endochrysea, P. MIN, rubropulchra

Physcia sciastra, P. ISC, KANU, Phaeophyscia sciastra Y 17 2013 ss, n lithotea MSC, MIN, MSC, WIS, Phaeophyscia squarrosa Physcia lacinulata 29 2006 MIN, KANU, Rinodina nimbosa, R. Phaeorrhiza nimbosa 1 1897 MIN phaeocarpa

Phlyctis boliviensis Y* 3 2013 ISC I,MT, e, ii, Physcia adscendens Y 7 2000 ISC, MIN n

ISC, KANU, MSC, OMA, Physcia aipolia Y 128 2013 I,tt,e,i-t,n NEB, MIN, WIS, ISC, MSC, Physcia americana Y 48 2013 MIN, e KANU ISC, NEB, Physcia caesia 10 2011 ss, ii, n MIN KANU, Physcia dakotensis 18 2007 TLE, MIN, KANU, Physcia halei 6 2002 MSC, MIN,

86 ISC, KANU,

MSC, TLE, Physcia millegrana Y 114 2013 l, n, ttt MIN, WIS, BRY, NEB

Physcia neogaea 3 2012 ISC

Physcia phaea Parmelia phaea Y 7 2013 ISC, KANU

Physcia pumilior 2 2011 ISC, KANU ASU, ISC, KANU, MSC, OMA, Physcia stellaris Y 262 2013 SBBG, TLE, I, ss, m, ii UPS, WIS, NEB, MIN, BRY, ISC, KANU, Physcia subtilis Y 16 2007 TLE, MIN, WIS

Physcia chloantha, P. ISC, KANU, luganensis, MSC, Physciella chloantha Y 76 2013 Phaeophyscia SBBG, chloantha MIN, Physcia melanchra, COE, ISC, Physciella melanchra Phaeophyscia Y 23 2013 KANU, melanchra MIN, ISC, MSC, Physconia detersa Physcia detersa Y 59 2013 I, tt MIN, WIS,

Physconia enteroxantha Y 3 2012 ISC, SBBG e,ss

ISC, DUKE, Physconia leucoleiptes Physconia kurokawae Y 65 2011 KANU, TLE, MIN,

Catapyrenium tuckermanii, Dermatocarpon tuckermanii, Dermatocarpon NEB, OMA, Placidium arboreum 8 1898 arboretum, MIN, WIS Endocarpon tuckermanii, Endopyrenium tuckermanii

ISC, MONT, Catapyrenium Placidium lachneum 20 1990 TLE, NEB, lachneum MIN

KANU, Catapyrenium MSC, OSC, squamulosum, Placidium squamulosum 21 2011 TENN, Dermatocarpon MIN, SRP, hepaticum auct. non WIS

ISC, KANU, MONT, Placopyrenium fuscellum Verrucaria fuscella 19 2000 MSC, MIN, NEB Placynthiella icmalea Saccomorpha icmalea 2 2003 KANU

CANL, ISC, MSC, Placynthium nigrum Pannaria nigra 20 2007 SBBG, WIS, NEB, BRY, MIN

Acarospora chlorophana, A. Pleopsidium chlorophanum 2 1896 MICH, NEB erythrophora, A. texana, A. weldensis

88 Polyblastia cupularis 1 1963 ISC

Teloschistes ILLS, ISC, Polycauliona candelaria candelarius, Xanthoria 25 1994 I, tt,e,i-t, n NEB candelaria Teloschistes polycarpus T. ISC, MSC, I, MT, e, Polycauliona polycarpa 38 1982 ramulosus, Xanthoria NEB, MIN s-I, n polycarpa, X. ramulosa Biatorella revertens, B. simplex, Lecanora ISU, KANU, Polysporina simplex 9 2002 privigna var. revertens, NEB, WIS Sarcogyne simplex Clathroporina nuculastrum, C. Porina nuculastrum confinis, 1 1961 ISC Polyblastiopsis floridana Huilia albocaerulescens, Porpidia albocaerulescens Lecidea Y 9 2013 ISC, MIN albocaerulescens, L. hebescens

ISC, ASU, Protoblastenia rupestris Y 9 2008 CANL, MSC, MIN

Psilolechia lucida Lecidea lucida 5 2011 ISC ISC, KANU, MONT, MSC, OMA, Psora decipiens Lecidea decipiens 53 2009 TLE, NEB, MIN, TENN, WIS, ASU Psora globifera Lecidea globifera 1 1992 MIN ISC, KANU, Psora pseudorussellii 11 2008 WIS, MSC, NEB

Psora rubiformis Lecidea rubiformis 3 1978 ISC

SD, MN, IL, 89

Psoraglaena c.f. dictyospora Macentina dictyospora Y* 1 2013 ISC NE, MO

ISC, KANU, MSC, OMA, Parmelia bolliana, P. Punctelia bolliana Y 157 2013 SBBG, US, frondifera WIS, BRY, MIN,

Parmelia borreri, P. ISC, NEB, Punctelia borreri 12 1978 pseudoborreri WIS Punctelia caseana 2 2012 ISC Parmelia graminicola, Punctelia graminicola 6 2000 ASU, MIN Parmelia semansiana

US, MIN, Punctelia hypoleucites Parmelia hypoleucites 14 1963 WIS

ISC, MIN, Punctelia perreticulata Parmelia perreticulata 3 1992 e, ii MSC

COE, ISC, KANU, MSC, OMA, Punctelia rudecta Parmelia rudecta Y 192 2013 I, ss, n, ttt WIS, NEB, BRY, MIN, SBBG ISC, KANU, Pyrenula pseudobufonia Y 25 2013 MSC, MIN, NY, WIS

Pyxine caesiopruinosa 1 1979 ISC ISC, MSC, Pyxine sorediata Y 19 2013 NEB, MIN, WIS,

ISC, KANU, Ramalina americana Ramalina fastigiata Y 54 2014 NEB, MIN, S WIS

ISC, MIN, 90 Ramalina intermedia 17 2011 UPS, WIS

Ramalina sinensis 1 1899 MIN Lecidea elabens, L. Ramboldia elabens melancheima, 2 1894 MIN Pyrrhospora elabens KANU, Rhizocarpon disporum 6 2002 MIN Rhizocarpon grande 1 1967 BRY Lecanora chrysoleuca, ISC, KANU, Rhizoplaca chrysoleuca 16 2007 ss L. rubina NEB, MIN

Lecanora Rhizoplaca melanopthalma 4 1978 ISC ss melanophthalma

KANU, Rhizoplaca subdiscrepans 13 2000 MIN, WIS

ISC, MONT, Rinodina bischoffii 13 2001 MSC, NEB, WIS, OMA, US, Rinodina cana Rinodina iowensis 3 1895 WIS Rinodina chrysomelaena 1 NA WIS KANU, Rinodina destituta Rinodina vezdae 2 2002 MIN Rinodina efforescens Y* 2 2013 ISC WI, MN Rinodina exigua 2 1894 ISC, WIS I Rinodina freyi Rinodina glauca Y 2 2013 ISC Rinodina oxydata 1 1895 MSC Rinodina pachysperma 5 2011 KANU

Rinodina papillata 1 1957 MSC S 91

ISC, KANU, Rinodina populicola Y 18 2013 MIN, WIS,

With Arthonia Rinodina pyrina Y 2 2002 KANU, ISU radiata # 873346

Rinodina siouxiana 1 2002 KANU ILLS, ISC, Caloplaca elegans, C. KANU, splendens, Placodium Rusavskia elegans 49 2008 MSC, NEB, e, tt elegans, Xanthoria MIN. OMA, elegans WIS Xanthoria Rusavskia sorediata sorediata,Caloplaca 1 NA WIS sorediata

Sarcogyne clavus Biatorella clavus 1 1895 MIN Biatorella hypophaea, Sarcogyne hypophaea 3 1962 ISC, MIN B. nannaria, B. plicata

ISC, KANU, Sarcogyne regularis Biatorella pruinosa 12 2006 MSC, MIN Sarcogyne similis f. convexa 1 NA WIS Sclerophora farinacea 3 1895 MIN ISC, KANU, Coniocybe nivea, C. MONT, Sclerophora nivea 12 1896 pallida MSC, NEB, NY, WIS ISC, ASU, KANU, Scoliciosporum chlorococcum Bacidia chlorococca Y 10 2012 I, n MSC, MIN, WIS, BRY

DUKE, ISC, Scytinium dactylinum Leptogium dactylinium 20 1978 MSC, MIN, WIS COE, ILLS,

ISC, MSC, 92

Scytinium lichenoides Leptogium lichenoides Y 41 2013 SBBG, o NEB, WIS, MIN, Collema plicatile, ISC, NEB, Scytinium plicatile 5 1895 Leptogium plicatile MIN Leptogium Scytinium subtile minutissium, L. 2 1965 MSC, WIS perminutum, L. subtile KANU, Solitaria chrysophthalma 10 2009 MIN, Solorina saccata 1 1956 ISC Spilonema revertens 2 2002 KANU Squamarina lentigera Lecanora lentigera 2 1982 ISC DUKE, ISC, Caloplaca subsoluta, KANU, Squamulea subsoluta Caloplaca irrubescens, Y 61 2013 MSC, MIN, C. modesta WIS

Staurothele areolata 9 2012 ISC, MIN Endocarpon Staurothele drummondii drummondii, E. 1 1961 MIN wilmsoides

Staurothele fissa 2 1894 ISC, NEB

ISC, MIN, Stictis urceolatum Conotrema urceolatum Y 5 1895 WIS, NY, NEB

Strangospora pinicola 1 2002 KANU MICH, MSC, US, Strigula americana Arthopyrenia tenuis 11 1967 WIS, MIN, NY Arthopyrenia affinis KANU, Strigula jamesii 4 2003 auct. MIN Arthopyrenia faginea,

Strigula stigmatella Porina cinerea, P. 1 2009 KANU 93

faginea ASU, KANU, Arthopyrenia Strigula submuriformis 26 2009 MSC, OMA, submuriformis SBBG, WIS, MIN, NY, ISC, KANU, Teloschistes chrysophthalmus Y 105 2014 NEB, MIN, TENN, WIS,

Tetramelas insignis Buellia insignis 2 1993 SBBG Thelidium minutulum Y* 1 2012 ISC MO Thelidium pyrenophorum 5 1908 MIN

Thelidium zwackhii 1 2003 KANU T. epilithellum, T. Thelocarpon laureri majusculum, T. 2 1959 ISC, MIN prasinellum

ISC, MSC, Thrombium epigaeum 12 1982 WIS, BRY, MIN MIN, NY, Thyrea confusa 9 1963 WIS Toninia aromatica 3 1978 ISC Trapelia involuta, Trapelia glebulosa Lecidea gregaria, L. 1 2012 ISC ornata WI, MO, Trapelia placodoides Y* 1 2013 ISC MN, IL Lecidea aeruginosa, L. Trapeliopsis flexuosa 1 2010 ISC a flexuosa ISC, KANU, Lecidea granulosa, L. SBBG, Trapeliopsis granulosa 12 2001 a quadricolor MIN, NY,

WIS 94

Lecidea viridescens, ISC, KANU, Trapeliopsis viridescens Y* 3 2013 WI Micarea viridescens NY ISC, US, Cetraria halei, C. Tuckermannopsis americana 16 1896 MIN, WIS, ciliaris var. halei NEB

Tuckermannopsis ciliaris Cetraria ciliaris 1 1896 US a, sss

Likely collected Umbilicaria mammulata Gyrophora dillenii 1 NA mid 1970's ISC, WIS, Usnea angulata 8 1895 NEB, MIN collected once MO, MN, Usnea hirta Y 2 2013 ISC, MIN S-I, o, ii 1895 SD, NE, WI Usnea longissima Ach. 1 1894 ISC ASU, ISC, Usnea mutabilis 7 1978 MIN

Usnea barbata var ISC, WIS, Usnea strigosa 8 1895 a florida MIN, Fuscopannaria leucophaea, Pannaria ISC, MIN, Vahliella leucophaea leucophaea, 4 1894 WIS Parmeliella microphylla ISC, KANU, MSC, SRP, MIN, MSC, Varicellaria velata Pertusaria velata 27 1967 MONT, TLE, NEB, WIS Variolaria ophthalmiza Pertsuaria ophthalmiza 1 1965 MIN Variospora velana Caloplaca velana 1 2009 ISC WI, MN, IL, Verrucaria calkinsiana Y* 4 2013 ISC 95 SD, NE, MO

Verrucaria finkiana 1 NA WIS With Porpidia Verrucaria furfuracea Y 2 2007 KANU, ISU albocaerulescens # 872183

Verrucaria glaucovirens 4 1996 ASU, MIN ISC, KANU, Verrucaria muralis Y 30 2013 OMA, NEB, MIN ISC, KANU, Verrucaria nigrescens Y 25 2013 NEB, MIN Verrucaria phloeophila 2 2003 KANU ISC, NEB, Verrucaria rupestris 6 1963 WIS Verrucaria viridula 2 1903 ISC KANU, Xanthocarpia crenulatella Caloplaca crenulatella 7 2009 MIN

ASU, ISC, KANU, MSC, OMA, Xanthocarpia feracissima Caloplaca feracissima Y 44 2013 SBBG, NEB, MIN, WIS, BRY,

Xanthocarpia tominii Caloplaca tominii 1 2006 ASU

ASU, CANL, DUKE, ISC, Xanthomendoza fallax Xanthoria fallax Y 105 2013 KANU, S-I, e, ss MSC, WIS, BRY, MIN, OMA,

CANL, ISC, Xanthomendoza fulva Xanthoria fulva Y 51 2013 e MIN

ISC, KANU, Xanthomendoza hasseana Xanthoria hasseana 54 2012 e 96 MIN, WIS,

ISC, KANU, ulophyllodes Xanthoria ulophyllodes Y 52 2012 Xanthomendoza MIN, OMA

ISC, KANU, Oxneria weberi, Xanthomendoza weberi 90 2012 NY, MIN, Xanthoria wetmorei OMA, Parmelia conspersa, P. Xanthoparmelia conspersa 1 1895 NEB isidiata Xanthoparmelia cumberlandia Parmelia cumberlandia 3 2012 ISC, MIN ss ISC, KANU, Xanthoparmelia hypofusca Parmelia tasmanica 4 2000 MIN Xanthoparmelia lineola Parmelia lineola 1 2000 KANU ISC, KANU, Xanthoparmelia mexicana Parmelia mexicana 11 2000 MIN KANU, Xanthoparmelia plittii Parmelia plittii 2 2000 MIN

TOTAL 465 150 7907