Preliminary Lichen Flora for Portage County, Wisconsin

By

Mary E. Pawlowski-Bartkowiak

A Thesis

Submitted in partial fulfillment of the requirements for the degree

MASTER OF SCIENCE

IN

NATURAL RESOURCES (FORESTRY)

College of Natural Resources

UNIVERSITY OF WISCONSIN

Stevens Point, Wisconsin

May 2013

APPROVED BY THE GRADUATE COMMITTEE OF:

______Dr. James Bennett, Department of Botany University of Wisconsin-Madison

______Dr. Robert Freckmann Emeritus Professor of Biology & Water Resources

______Dr. Virginia Freire Associate Professor of Biology – Committee Co-Chair

______Dr. Richard Hauer Associate Professor of Forestry – Committee Co-Chair

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ABSTRACT

Many studies, both vegetation and floristic, have been conducted throughout

Wisconsin and have contributed to our knowledge of lichens in this state (Culberson

1955, Hale 1955, Foote 1966, Newberry 1974, Will-Wolf 1980, Bennett 2006a).

However, none of these studies have focused on Portage County, and cannot be used to represent the assemblage of lichens at present. Portage County lies in the center of

Wisconsin and offers a variety of interesting physiographic characteristics. The tension zone (Curtis 1959) traverses the county from east to west, suggesting that an overlap of northern and southern vascular plant species may occur in the area. The last glacial event provided Portage County with rolling terminal moraines, wetlands and sandy outwashes.

The results of a preliminary floristic study of lichens in Portage County,

Wisconsin are presented here. A total of 180 species in 73 genera were identified from field collections made in 2010 through 2012. A catalogue of species is included with 208 lichens presented as a result of field work, a literature search for previously reported collections and a physical search of the Wisconsin State Herbarium (WIS). There are

115 new records for the county and the following 3 species are reported for the first time for the State of Wisconsin: Leptogium rivulare, Heterodermia obscurata, and Peltigera extenuata. In addition to state and county records, two lichens of significant ecological interest were collected during this study; Lobaria pulmonaria, and Normandina pulchella.

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ACKOWLEDGEMENTS

I would like to thank the members of my graduate committee, Dr. Jim Bennett,

Dr. Bob Freckmann, Dr. Virginia Freire, and Dr. Rich Hauer for all of their support.

Each member provided me with consultation, direction and encouragement when I needed it the most. Thank you, to both the UWSP Freckmann Herbarium and the

Wisconsin State Herbarium-Madison, for allowing me area to work and access to the collections. I’d also like to thank Richard Olson, Ed Damask, and Quinton and Shari

Cieslewisz for permission to sample on their properties. Financial support was provided by the College of Letters and Science and the Botanical Club of Wisconsin, for which I am very grateful. Much appreciation goes to Ezekiel Behnke for assistance with herbarium work and Carol Kropidlowski for accompanying me in the field. I am also thankful for my academic family, there are too many individuals to name that have provided inspiration and support. Finally I would like to thank Rick and Ben for supporting me in this project.

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TABLE OF CONTENTS

ABSTRACT……………………………………………………………………………...iii

ACKNOWLEDGMENTS………………………………………………………………..iv

LIST OF TABLES………………………………………………………………………..vi

LIST OF FIGURES……………………………………………………………………...vii

INTRODUCTION AND LITERATURE REVIEW……………………….……………..1

General Background and Biology…………………………………………………2

Ecological Importance…………………………………………………………….7

Environmental Indicators/Monitoring With Lichens……………………………...9

Human Interactions………………………………………………………………12

Animal Interactions………………………………………………………………14

Wisconsin Lichens……………………………………………………………….15

Significance of a County Flora…………………………………………………..17

METHODS AND MATERIALS...... ………………………………………………….19

Study Area……………………………………………………………………….19

Field and Lab Methods…………………………………………………………..22

RESULTS/DISCUSSION…………………………………………….…………………28

Rare and Significant Lichens…………………………………………………….29

Culberson Site/Curtis Stand 3044 Revisited…………………………………….34

Previous Reports…………………………………………………………………38

COLLECTION SITES…………………………………………………………………...40

CATALOG OF SPECIES………………………………………………………………..47

LITERATURE CITED…………………………………………………………………..59

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LIST OF TABLES

Table 1. Percent cover by habitat type for Portage County...... 24

Table 2. Top 18 lichen genera collected from 2010 – 2012……………………….29

Table 3. Lichen species identified for Culberson site……………………………...37

Table 4. Lichens previously reported for Portage County………………………… 39

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LIST OF FIGURES

Figure 1. Map of Wisconsin with Portage County highlighted…………………….17

Figure 2. Surface and subsurface features of Portage County……………………..18

Figure 3. Map of Wisconsin with tension zone transecting Portage County……… 19

Figure 4. Map of study sites in Portage County……………………………………20

Figure 5. Map of Portage County landcover. ………………………………………22

Figure 6. Habitat of Leptogium rivulare on the Tomorrow River………………….27

Figure 7. Google Earth image of Cieslewicz property…………………………….30

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INTRODUCTION

Lichens are often mistaken for moss and referred to as plant-like organisms.

Although they are of small stature, they are neither simple nor a single organism but a composite represented by two and sometimes three biological kingdoms. The delicate balance that is required for this symbiosis to remain successful is dependent on many complex interactions. It has been well documented that lichens are excellent indicators of air quality and have been used historically to monitor for pollutants (Hawksworth & Rose

1970, Newberry 1974, Will-Wolf 1980, Richardson 1992, Gries 1996).

Many studies, both vegetation and floristic, have been conducted throughout

Wisconsin and have contributed to our knowledge of lichens in this state (Culberson

1955, Hale 1955, Foote 1966, Newberry 1974, Will-Wolf 1980, Bennett 2006a).

However, none of these studies have focused on Portage County, and cannot be used to represent the assemblage of lichens at present. Portage County lies in the center of

Wisconsin and offers a variety of interesting physiographic characteristics. The tension zone (Curtis 1959) traverses the county from east to west, suggesting that an overlap of northern and southern vascular plant species may occur in the area. The last glacial event provided Portage County with rolling terminal moraines, wetlands and sandy outwashes.

Results of a preliminary floristic study of lichens in Portage County, Wisconsin are presented here. 208 lichen species are presented with 115 reported as new records for the county and 3 new records for the State of Wisconsin.

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General background and biology of lichens

Lichens are a composite life form consisting of a fungus (mycobiont) and an organism capable of producing carbohydrates through photosynthesis. Recent literature often refers to lichens as miniature ecosystems and not as organisms at all (Hinds &

Hinds 2007). The photosynthetic symbiont, known as a photobiont can be an alga or cyanobacteria, or occasionally both. Three biological kingdoms may comprise the lichen when all three are present; Fungi, Protista and Monera. The tripartite association involves the fungus with a green alga as a primary photobiont and the cyanobacteria in distinct packets known as cephalodia. The mycobiont is most often from the

Ascomycetes group of fungi which are characterized by the production of spores in sac like structures. Occasionally the mycobiont is from the Basidiomycetes group which represents the mushroom producing fungi. The photobiont is capable of producing food for the partnership and the mycobiont provides protection, ambient light and moisture for the association. A simplistic overview: the mycobiont provides structural support in the form of a thallus and the photobiont is the occupant that produces food for both associates.

The lichen symbiosis is very complex and research continues on the subject.

Lichens were previously believed to be an example of mutualism, i.e. with all partners benefiting from the association. However, the union may be better described as varying degrees of parasitism (Nash 1996). In order for the mycobiont to access the food produced, it must invade the photobiont. This invasion suggests that the mycobiont may be benefiting more from the association than the photobiont. When the two components are grown separately in the lab, the mycobiont does not develop into a structure

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resembling a lichen thallus. Instead it forms a gelatinous mass. In contrast, the photobiont can be grown separately but does not look the same as when it is in the symbiotic state. The most common genus of green algae to appear in the lichen association is Trebouxia (Brodo 2001), which is rarely if ever found free living in nature.

Trentepohlia, another common photobiont, is able to form filamentous free living colonies. Cyanobacteria are commonly found free living; the two most common are

Nostoc and Gloeocapsa (Brodo et al. 2001). These photobionts, while in the lichen association do not reproduce or develop filaments. However, they are able to colonize extreme environments that they would not be able to live in outside of the association

(Richardson 1974).

More than 700 secondary compounds, contributing up to 20% dry weight, have been identified from lichens (Purvis 2000). The compounds produced are very diverse; the most common are the phenolic acids, the depsides and depsidones (Brodo et al.

2001). These substances appear to be concentrated in the thallus cortex with several possible functions: UV light screen, antibiotic and antifungal properties and allelopathic chemicals. There are many theories regarding the possible functions or strategies employed by the lichen in regards to these substances, however, it is agreed that they are not merely a waste product.

The thallus represents the vegetative body of the lichen and is mainly made up of the mycobiont rather than the photobiont. The four general growth forms that describe the various lichen thallus types are: foliose, fruticose, crustose, and squamulose. Other intermediate growth forms exist that are combinations or gradients of these main types.

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Although lichens are occasionally grouped together by thallus shapes for identification purposes, this does not imply that similar shaped lichens are related (Brodo 2001).

Foliose lichens are flattened, leaf like and are composed of distinct layers

(stratified). The upper cortex serves as an outer surface, made of tightly packed fungal hyphae. The medulla lies in the middle of the thallus; this is an area of loosely entwined hyphae. A layer of algae lies within the upper medulla, just below the upper cortex, wrapped by the hyphae. Occasionally there is a lower cortex which is also an area of tightly packed hyphae. If no lower cortex exists, the fungal hypha from the medulla is visible from below. Foliose lichens often have special structures (rhizines or haptors) to attach either loosely or firmly to the substrate on which they are growing. Although they resemble roots of vascular plants they do not absorb or conduct water or minerals.

Fruticose lichens differ from the foliose in that they do not have a defined upper and lower surface and they are three rather than two dimensional. Fruticose includes upright, shrubby, pendant, tufted, erect and branched growth forms. The branches of these lichens can be round or somewhat flattened and attached at a single point or at multiple locales.

Crustose or crust lichens are firmly attached and sometimes embedded with the substrate on which they are growing. They have no lower cortex and are almost impossible to remove from the growing substrate. Therefore, they need to be collected with the substrate for identification purposes (Brodo et al. 2001). Of these main growth forms, the crustose are the most difficult to identify; experienced lichenologists concur that some are impossible to identify in the field (Bennett & Wetmore 2004).

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Squamulose lichens are best described as a combination of a crust and foliose growth form. The thallus is comprised of small overlapping scales which are attached to the substrate on one edge. Squamulose and crustose growth forms are known collectively as microlichens; foliose and fruticose are referred to as macrolichens.

All forms exhibit slow growth rates, with the crustose lichens being the slowest at one millimeter or less per year (Richardson 1992). Unimpaired by this slow growth rate, crustose lichens colonizing rocks are able to penetrate several millimeters into a rock substrate with their fungal hyphae. The secondary compounds produced by the lichen expedite the weathering process and help to initiate the first step in soil formation from rock. Although other organisms are capable of colonizing bare rock, lichens represent the most abundant and diverse group of invaders (Brodo et al. 2001).

Lichens are capable of vegetative reproduction via specialized propagules that contain both the mycobiont and the photobiont. Soredia are powdery assemblages of fungal hyphae surrounding photobiont cells and are released from the photobiont layer through gaps of the thallus cortex known as soralia. Some lichens are completely engulfed in soredia and are then described as leprose. The location of the soredia on the thallus is important for identification purposes. Isidia are similar vegetative propagules in that they also contain both the mycobiont and the photobiont, however, they possess a smooth outer cortex and appear as outgrowths of the thallus. Both isidia and soredia are easily separated from the thallus and dispersed by water, wind or animals and if proper conditions exist, they will form a new lichen.

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The fruiting body of a lichen is expressed by the mycobiont. Since the majority of lichen fungi belong to the Ascomycetes, they produce spores in ascomata and are of two basic types: apothecia and perithecia. Apothecia are linear, cup or disc shaped with an exposed surface for spore dispersal. There are different forms of apothecia, but basically there are two types, those that contain algae in the margin and those that do not.

These specialized forms of apothecia are important for classifying lichens. Perithecia are flask shaped and embedded in the lichen thallus, with spores exiting through a small hole known as an ostiole. Lichen fruiting bodies are long lived and perennial, but can be preyed upon by a number of invertebrates (Brodo et al. 2001).

As soil type is important to vascular plants, substrate is of equal importance to lichens since they are frequently in intimate contact. Identification keys often employ substrate types in their couplets since lichens have preferences for specific surfaces. The main categories of substrates include: saxicolous (rock dwelling), terricolous (ground), corticolous (bark), lignicolous (wood). The most important substrate criteria for lichen habitat include chemistry (pH), texture and moisture (Culberson 1955; Brodo 1973). In

Wisconsin, the majority of lichens occur on woody substrates and are corticolous or lignicolous (Will-Wolf in Waller 2008). The favored rock substrates for saxicolous lichens are those rich in calcium carbonate (lime). Completely different lichen communities will inhabit the calcareous substrates than the siliceous granites or schists

(Brodo et al. 2001).

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Ecological Importance

Lichens are very successful at pioneering harsh environments; they are often able to inhabit areas where neither symbiont could survive alone. Strategies employed by lichens include: drought tolerance, ability to photosynthesize and make their own food, utilize small amounts of nutrients that land on thallus as dust, and produce very small propagules (Brodo 2001). Cyanolichens (lichens with cyanobacteria as one of the symbiotic partners) are able to fix atmospheric nitrogen and comprise approximately 10% of lichen species. Several lichens contain both green algae and cyanobacteria as the photobiont, which provides them with an additional advantage. The two photobionts have different moisture requirements for net photosynthesis: green algae are able to photosynthesize with the presence of water vapor and cyanobacteria require liquid water

(Friedl & Budel 1996). A lichen thallus that includes both of these photobionts will be more adaptable to a wide variety of environments and climatic variations.

Lichens accumulate nutrients throughout their life and release them into the ecosystem as they decompose. As expected, cyanolichens contribute more nitrogen than lichens with green algae as their photobiont. Epiphytic lichens that live lower in the canopy receive nutrients that are leached from the higher canopy. Pike (1978) found that nitrogen was the most significant nutrient contribution made by epiphytic lichens, followed by phosphorous and potassium, with even less amounts of calcium and magnesium. These mineral contributions were minimal compared to other biomass contributors such as coarse and fine woody debris and leaf fall.

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Weathering of rocks to form soil is often initiated by lichens as their fungal hyphae penetrate the rock up to several millimeters in depth. Acids formed through regular lichen metabolism and secreted through the hyphae in the rock also aid in the weathering process (Purvis 2000). As saxicolous lichens die, they form organic matter on the rock surface. This matter traps other particles including spores and seeds which in turn continue to accumulate on the surface and small areas of vegetation begin to pioneer a new community (Brodo et al. 2001).

On fertile soils, lichens cannot compete with the speed in which vascular plants and bryophytes are able to colonize. However, on disturbed and sandy soils that are too harsh for many plants, lichens are able to establish a community and make significant ecological contributions. Such services/contributions include: soil stabilization, soil nutrients/organic matter, moisture retention, invertebrate habitat, albedo and seed entrapment.

Some lichens have exhibited allelopathic tendencies, in that they excrete enzymes that may prevent the germination of some seeds (Brodo et al. 2001). On the contrary,

Opuntia fragilis growing in Wisconsin appeared to be benefiting from growing in close proximity with two lichens; Cladonia mitis, C. rangiferina and a spikemoss; Selaginella rupestris. However, this could be another example of allelopathy, to which cactus is resistant to. The lichens may be preventing other plants from growing near and competing with the cacti for moisture and nutrients (Bennett et al. 2003).

Icmadophila ericetorum has been reported as having parasitic relationship with bryophytes by Richardson (1974). The fungal hyphae of the lichen have been found

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penetrating dead stems of the moss Mnium hornum on an old stump in Scotland.

Thelotrema subtile has allelopathic tendencies, inhibiting the growth of the small brown liverwort Frullania (Richardson 1974).

Environmental Indicators/Monitoring With Lichens

Lichens rely almost entirely on the atmosphere for nutrients. Unlike vascular plants, they lack a protective cuticle; therefore impurities can concentrate on the thallus and spread easily into the photobiont layer, causing mortality (Gries 1996). This sensitivity makes lichens efficient indicators of air quality, which is why they are used in biomonitoring programs. Sensitive lichens are intolerant of sulfur dioxide (SO2) and ammonia (NH3) (Richardson 1992) and these two pollutants are of local concern in

Portage County. Coal burning power plants and pulp paper mills are major contributors of sulfur dioxide in Wisconsin (Newberry 1974; Will-Wolf 1980) and have lethal effects on lichens sensitive to acids. Other industries and fossil fueled combustion engines also contribute sulfur dioxide into the atmosphere. Ammonia and nitrogen deposition from fertilizers and livestock change the pH of surrounding substrates, eliminating sensitive lichens and favoring nitrophilous lichens (Jovan 2005). Lichens preferring acidic environments are acidophiles and those preferring more basic pH are nitrophiles.

Because of their larger surface area, fruticose and foliose lichens are more sensitive to air pollution than crustose lichens and are most often used for air quality monitoring projects

(Pinho 2004).

Shifts in lichen communities occur with pollution gradients. Hawksworth and

Rose (1970) illustrated how specific lichens have different sensitivities to sulfur dioxide

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levels. Zones of lichen communities are identified radiating from point source pollution; diversity, abundance and cover increases with distance. Epiphytic lichens are the most sensitive to atmospheric impurities (Richardson 1992). On the other hand, there are lichens that require impure air. The crustose lichen, Lecanora conizaeoides, is an obligate acidophile native to Europe, and is indicative of polluted air quality. This lichen spread throughout Europe following increased levels of sulfur dioxide during the early

20th century and has now arrived in North America (LaGreca 2006). Pollutant tolerant lichen species include Candelaria concolor, Phaeophyscia spp., Physcia spp. and

Xanthoria spp. (Jovan 2005). While investigating the effects of a pulp paper mill on corticolous lichens in central Wisconsin, Newberrry (1974) found Scoliciosporum chlorococca to be one of the most pollution tolerant species in the area. Four other lichens found to be tolerant were: Lepraria spp., Flavoparmelia caperata, Punctelia rudecta and Physcia millegrana, correlating with results of other pollution monitoring surveys (Nash 1972). Physcia millegrana, the most pollution tolerant macrolichen in eastern North America, is the only macrolichen found within New York City’s Central

Park (McCune 2000).

Lichens are recognized as indicators of forest health by the U.S. Department of

Agriculture Forest Service (USDA FS). Beginning in 1990, macrolichens were used in the Environmental Monitoring and Assessment Program, followed by the Forest Health

Monitoring (FHM) Program from 1993-1999. Commencing in 2000, lichen indicators have been utilized in the Forest Inventory and Analysis (FIA) Program. The objective of the FIA program is to perform surveys on permanent plots throughout the United States and to monitor trends over time. Lichen sensitivity makes them efficient indicators of air

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quality that could potentially impact the overall health of a forest ecosystem. At present, three major categories have been outlined for lichen indicators: air quality, climate and biodiversity. It is anticipated that other categories will be added in the future and may require investigation into the effects of surrounding land use on the forests (Will-Wolf

2010).

The richest distribution of lichens across habitat types is often found in old- growth forests. Microhabitats increase with age within a forest, providing habitat for a wide range of lichen species (McMullin et al. 2008). Because of this, much research has been focused on identifying old growth or ancient forests by the suite of lichens that are present. Rose (1976) observed that certain epiphytic lichens were indicative of old growth forests in Britain. He developed an Index of Ecological Continuity (IEC) from a list of 30 lichens. This index was used to measure the continuity of the stand, which could thereby verify the old growth status. Ongoing research has included lists of lichen species for angiosperm and gymnosperm dominated forests (Selva 1994). The reasons why some lichens are restricted to old growth stands could be due to the following: these species grow very slow, they have very specific habitat requirements (often based on temporal elements), and lack efficient means of dispersal (Hinds & Hinds 2007).

Cyanolichens are the most sensitive to air quality and other disturbances

(Cameron & Richardson 2006). They comprise about 10% of all lichen species and because they require liquid water for photosynthesis (Lange et al. 1986) they are often found near water or in areas with high humidity. Epiphytic cyanolichens are especially sensitive to air pollution (Gries 1996). Ironically, free living cyanobacteria are extremely adaptable and able to tolerate great shifts in temperature, light and salinity. Acidity is

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the largest limiting factor, as cyanobacteria are unable to inhabit acidic environments; preferring a pH greater than 7.

Since lichens are an assemblage of different organisms behaving as a single entity, it is important to remember that each symbiont may relate differently to the same disturbance. Any shift in the balance between the mycobiont and photobiont(s) could destroy the partnership. The effects of urbanization are complex and changes to lichen communities are not often easy to identify (Gries 1996).

Human Interactions

Rock tripes or ‘tripes de roches’, species of Umbilicaria and Lasallia were historically used as a food source by native people and occasionally by Arctic explorers who found themselves with limited rations. According to John W. Thomson (2003), the earliest reference to lichens in Wisconsin was from the Jesuit Relations of 1663-1664 where “Tripe de Roche” was consumed in central Wisconsin, enroute to Lake Superior’s

Chequamegon Bay during October 1660. This statement refers to Umbilicaria species.

Lecanora esculenta, a squamulose lichen found in arid areas of the Middle East has also been used as a food by humans. Small patches of the lichen fall off the rocks following heavy dew. The lichen is then collected, added to meal and made into bread. It is suggested that this is the legendary lichen that fed the Israelites during their exodus from

Egypt (Richardson 1974). Several other lichens have been used as a food source throughout history, these include: Bryoria spp., Cladonia spp., and Cetraria spp. (Brodo et al. 2001).

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Lichens contain antibiotic substances that are effective against gram positive bacteria (Burkholder et al. 1944). Vartia (1973) reviewed and summarized the antibiotic reports of lichens and concluded that more than 50% of the lichens tested have antibiotic properties. Usnic acid, responsible for the pale yellow or ‘lichen yellow’ of many lichens, is easily recognized and is one of the most effective of the lichen antibiotics.

Various species of Bryoria, Usnea and Alectoria exhibit both antibacterial and absorptive properties. Flavoparmelia caperata, Lobaria retigera, Parmelia saxatilis, Parmotrema perforatum, P. perlatum and Peltigera aphthosa are a few lichens that have been used medicinally as tonics, laxatives, expectorants, or topically as salves (Brodo et al. 2001).

Cetraria islandica, known as “Iceland moss” has been used for centuries in Iceland for a variety of illnesses (Podterob 2008). Hundreds of unique organic compounds are produced by lichens; as molecular work continues, more will be revealed. It seems reasonable to expect that new uses for these compounds will be discovered as well

(Purvis 2000).

Historically, lichens have been an important source for dyes throughout the world.

The sources of colors in lichens are the same as the substances used for identification purposes (Brodo et al. 2001). Production of cudbear (purple dye) in 18th century Europe nearly decimated the crustose lichen Ochrolechia tartarea by over harvesting (Casselman

2001). The popularity of the Harris Tweed was due not only to the color of the weave but to the moth repellency of the fabric as well. The lichen provides the wool with a bitter taste that repels the larvae of the moths (Richardson 1974). Those that provide the deep purple and red dyes contain the lichens substance erythrin in addition to lecanoric,

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gyrophoric and alectoronic acids and have a positive red reaction when bleach is applied to the medulla (Brodo et al. 2001).

Animal Interactions

It is widely accepted that a variety of lichens including several species of

Cladonia provide winter food for caribou (Rangifer tarandus) in North America. A substantial amount of their seasonal diet is comprised of terricolous, saxicolous and corticolous macrolichens (Richardson & Young 1977). White-tailed deer (Odocoileus virginianus) also feed on lichens in winter, utilizing both Evernia mesomorpha and

Usnea spp. below a visible browse line (Hodgman & Bowyer 1985). Lichens are high in carbohydrates yet low in proteins (Richardson 1974). Ironically, lichens that fix atmospheric nitrogen and would therefore contain proteins are not typically browsed

(Brodo et al. 2001). Flying squirrels (Glaucomys sabrinus) and red-backed voles

(Clethrionomys gapperi) of the Northern Rocky Mountains also have low nutrient diets comprised mainly of lichens (Alectoria spp. and Bryoria spp.) and fungi (Dubay et al.

2008). In addition to food, nest boxes of flying squirrels were found to be constructed with over 90% Bryoria species (Hayward & Rosentreter 1994).

Lichens provide food, camouflage and oviposition sites for invertebrates.

Orabatid mites, springtails, psocids, rotifers, tardigrades, snails and slugs are known predators of lichens. Stubbs (1989) found a positive correlation between corticolous lichens and several species of invertebrates on Quercus rubra in Maine. Not all lichens are predated equally, those with a more developed set of secondary compounds tend to be avoided (Lawrey 1989).

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In Europe, a night flying moth, Biston betularia, was studied for its versatile appearance. Since the moth was immobile during the day, the ability to camouflage itself is a high priority for survival. While living with abundant lichen flora, the moth exhibits mottled shades of grey, brown and white. This allows the moth to blend in with the surrounding lichens and avoid becoming prey to birds. As air quality plummeted, and lichens disappeared from trees, the moth had evolved to a darker more melanin appearance; this solid morph blends in with the trees that are void of lichens (Bishop et al. 1975). This lichen/moth interaction demonstrates evolution within a relative short time period. Although the melanic morph form is complicated and as expected there are many other factors besides air quality that have impacted the physical appearance of this moth, it continues to react to the environment (Cook 2003). As the boles of trees lighten in response to reduced soot emissions the mottled appearance of the moth has returned.

Recent research has revealed that three lichens (Parmelia sulcata, Cladonia rangiferina and Lobaria pulmonaria), all native to Wisconsin, contain an enzyme capable of breaking down prion proteins responsible for chronic wasting disease. Chronic wasting disease is an infectious disease of white-tailed deer and has been known to occur in deer farms throughout Wisconsin (Johnson et al. 2011).

Wisconsin Lichens

Lichenology in Wisconsin is deeply rooted, with Increase A. Lapham and Thomas

Hale collecting lichen specimens from 1858 – 1860 (Thomson 2003). Toward the turn of the 19th century, Lellen L. Cheney collected lichens throughout the state. His canoe trip down the Wisconsin River during the summers 1893 and 1894 provided a sample of

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lichens growing throughout the state from Lac Vieux Desert to Prairie du Chien. This was accomplished prior to the presence of pulp paper mills, coal burning power plants and major dams that presently obstruct the river. Included in his samples, that were discovered in the WIS herbarium over 100 years later, were 25 lichen specimens collected from Portage County (Thomson 1998).

In 1944 John Thomson joined the Botany Department at the University of

Wisconsin-Madison; he contributed vastly to the knowledge of lichens through the state and was instrumental in guiding research opportunities with graduate students. In 2003, he published Lichens of Wisconsin, a guide to more than 600 species of lichens in the state. In 1965, the American Bryological Society held a foray in Wisconsin, and reported an additional 234 species for the state (Brodo 1967). During spring of 2002, a group of

Tuckerman Workshop lichenologists gathered in northern Wisconsin and spent five days exploring the lichen communities in three northern counties; they reported 130 new county records and 47 new state records (Lay 2004). Several ecological and floristic studies have been completed throughout the state but none have focused on Portage

County (Culberson 1955, Hale 1955, Will-Wolf 1980, Foote 1966, Newberry 1974,

Bennett & Wetmore 2009).

William Culberson (1955), studied corticolous lichens in upland forests in northern Wisconsin and one of his research sites was located in northern Portage County.

Field notes and a list of lichens identified for the site were located in the Plant Ecology

Lab (PEL) in UW-Madison. The site was revisited in 2012 for this study.

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Much of the research that has been completed in Wisconsin was based on ecological surveys that did not include searching for lichens in unique micro habitats.

Ecologically based surveys with random sampling do not always include the special habitats where rare lichens are found. Concurring with Bennett, Brodo (1967) reiterates that even in well studied areas, new records are often found following intense sampling.

The catalogue of lichen species included with this report provides an updated list for the county.

Significance of a County Flora

A flora is an inventory of species present within a specific geographical region. A floristic survey offers no quantitative data, and each species listed contributes equally to the compilation of species. County floras are valuable for many reasons but it is important to note that they are never complete. As natural and anthropogenic disturbances occur across the landscape, lichen genera and assemblages are in a constant state of flux.

A thorough investigation of an area can detect unique and delicate ecological communities based on the occurrence of sensitive lichen species. The identification of these areas may aid in conservation programs to protect biodiversity and assist natural resource management in land use planning. A flora of a county can also be used as a reference for future comparisons and may also conjure interest in conservation of the natural world. An initial list provides the base on which forthcoming surveys are able to build upon as well as a baseline for future comparisons. Voucher specimens are useful to

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note geographic distribution and may also be utilized by lichenologists to study taxonomic variations.

“What a thousand acres of Silphium looked like when they tickled the bellies of the buffalo is a question never again to be answered, and perhaps not even asked.” ~Aldo Leopold

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METHODS AND MATERIALS

Study Area

Portage County is located in the geographic center of Wisconsin (latitude 44° 29’

11”N, longitude 89° 31’20”W) and covers approximately 208,753 hectares (Figure 1).

The county is relatively flat with an average elevation of 335 m above sea level. Average annual rainfall is 81 cm, and average annual snowfall is 104 cm. There are 136 lakes and

7 rivers; the Wisconsin River serves as the major watershed.

Figure 1. Map of Wisconsin showing Portage County.

Three distinct surface and subsurface features are present within the county: residual area, moraines, and sand plain (Figure 2- Tesch 1982). A series of lateral moraines occur in the glaciated eastern part of the county and consist of loamy soils developed from glacial outwash. The residual area is present in the northwest portion of the county, bounded by the Wisconsin River to the south and the Plover River to the east.

The residual soils that occur in this area have formed in place rather than being deposited

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by the glaciers and are loamy and silty, covering igneous and sandstone bedrock.

Precambrian igneous rocks are exposed along the Wisconsin River with occasional outcropping within the northern portion of the residual area. Cambrian sandstone is also exposed in a few locations. The sand plain occurs between the moraines and the residual area and was once covered by Glacial Lake Wisconsin. Deep sandy and acidic soils are evident throughout the southwest corner of the county. Sand blows and Cambrian sandstone outcroppings are present within the sand plain. Granitic glacial relicts occur throughout the northeast portion of the county. Limestone may be found only as glacial drift from the Niagara Escarpment; no outcroppings are known to exist.

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Portage County lies within the tension zone as described by Curtis (1959). This ecotonal region, about 20-40 km wide, traverses the county from east to west, suggesting that an overlap of northern and southern vascular plant species occur in the area (Figure

3). Northern plant communities that reach their most southern boundary are the mixed conifer-hardwood stands of sugar maple (Acer saccharum), white pine (Pinus strobus), hemlock (Tsuga canadensis) and yellow birch (Betula alleghaniensis), and Sphagnum bogs of spruce (Picea spp.) and tamarack (Larix laricina). Prairies, pine and oak savannas, and hardwood forests of the southern province reach their northern limit within the zone. Curtis mentions that the following plant communities exist in Portage County:

Forests - southern and northern lowland, northern mesic and xeric; sand barrens and bracken grasslands; savannas, shrub-carr, alder thicket, fens, sedge meadows, and bogs.

Figure 3. Tension zone transecting Portage County.

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Field and lab methods

Lichens were collected beginning in 2010 and continued throughout 2012.

Approximately 700 specimens were collected at 58 localities throughout the county.

Several biologically diverse locations were sampled multiple times. A list of collection sites with applicable collection numbers appear in the Collection Sites section on page 40 and are indicated on Figure 4. Locations were chosen to include a full range of habitat types and plant communities, or simply anywhere that lichens were found within Portage

County. An effort was also made to find locations that would be rich with lichens. The following habitats were studied: forested areas, sand barrens, conifer bogs and swamps, sandblows, rocks and rock outcroppings, grasslands, roadsides, savannas, streams and river banks. Man-made substrates such as old foundations, fences, roofs, metals, and cemeteries were also searched. Special attention was paid to microhabitats within each sampling area to find the rare or less abundant species. Microhabitats included, but were not limited to: standing dead trees (snags) and down logs at varying degrees of decomposition, moss covered boles of trees, fence rails and posts, road-cuts, and partially submerged substrates at water edges.

An estimate of percent cover by habitat type (Table 1) was completed for the county using ARC-GIS software and the National Landcover Database from 2001

(Figure 5). Forested land is the most abundant cover type within Portage County. Barren land comprised of exposed rock, sand, & clay is represented by less than 1%.

Accordingly, most sampling sites were from forested areas with the least sampled from barren areas. Discrepancies occurred in woody wetlands and crop lands. No lichens

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were collected from crop lands and significantly more than 17% of the sampling sites were from woody wetlands. Multiple habitat types were present at many locations.

Figure 4. Study sites in Portage County WI, indicated by black filled circles.

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Table 1. Percent cover by habitat type for Portage County based on National Landcover Database 2001. Number of sampling sites with percent total for project. Number of Percent Cover Portage Habitat Type Sampling Total County Sites Forested 55.5% 44 38.3%

Developed 12.8% 15 13.0%

Woody Wetlands, Open Water, 16.9% 41 35.7% & Emergent Wetlands Crops, Pasture, Hay 8.6% 0 0.0%

Shrub, Scrub & Grasslands 6.1% 12 10.4%

Barren (rock, sand, & clay) 0.1% 3 2.6%

Total 100 115 100

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Figure 5. Map of Portage County landcover utilizing National Landcover Database 2001.

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Equipment used to collect lichens in the field included hand lens, knives, wood chisel, carbide tip chisel, hammer, distilled water, paint scraper, camera, and field notebook. Specimens were collected and placed in brown paper bags in the field.

Latitude, longitude and elevation were recorded using a Garmin GPSMap 60Cx unit utilizing the map datum WGS 84.

To eliminate live invertebrates from the samples, all specimens were frozen for 24 hours before being packeted into temporary collection envelopes to await identification.

Lichens were identified in the lab with various chemical tests and the use of a Nikon

Eclipse E400 compound microscope and an Olympus SZX12 dissecting microscope.

Following confirmation of identification, the specimens were re-packeted in 100% cotton paper with appropriate label data.

Voucher specimens are deposited in the University of Wisconsin–Stevens Point

(UWSP) Freckmann Herbarium. Label data for all specimens will be entered into the

Consortium of North American Lichen Herbaria (CNALH) database http://lichenportal.org. A representative set of duplicates will be sent to the WIS herbarium. Nomenclature follows Esslinger’s North American Lichen Checklist (2012).

Taxonomic keys used for identification purposes include: Brodo et al. (2001), Brodo

(2011), Hinds & Hinds (2007), Thomson (2003) and a variety of monographs.

Specimens were identified using chemical spot tests, and microscopic techniques described in Brodo et al. (2001). Specimens were checked by Dr. James Bennett at UW

Madison.

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To determine if new, rare or significant lichens were discovered, a search of published literature was performed to create a list of lichens known to exist in Portage

County prior to this study. The list was reviewed and updated to reflect present nomenclature. The WIS herbarium was searched for unpublished lichen specimens collected in Portage County.

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RESULTS/DISCUSSION

During this investigation from 2010 through 2012, approximately 700 specimens were collected from 58 study sites in Portage County. Some 603 specimens have been identified with the remainder either requiring further study or were not identifiable. A total of 180 species in 73 genera were recognized with the proportional following life forms: 34% crustose, 44% foliose, 21% fruticose and 1% squamulose. The majority of unidentified specimens are crustose. These are the most difficult of the life forms to identify and often require thin layer chromatography (TLC) for proper identification. At this time, resources were not available to perform such tests. In agreement with Will-

Wolf (in Waller 2008), the majority of lichens collected were from woody substrates, the percent by substrate is as follows: 66% corticolous, 6% lignicolous, 16% saxicolous, and

12% terricolous. The search of the WIS herbarium revealed four new reports for lichens in Portage County; Aspicilia cinerea, Candelariella vitellina, Fuscidea recensa, and

Pertusaria macounii.

The genera with the most species occurring throughout the county are shown in

Table 2. The fruticose lichen genera Cladonia is well represented by 30 species, which is not surprising as it is one of the largest genera in North America with more than 170 species. Of the 18 genera 50% (9 genera) are foliose and the remaining 9 genera are crustose (5 genera) and fruticose (4 genera).

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Table 2. Number of species in the top 18 lichen genera collected from 2010 – 2012. Cladonia 30 Aspicilia 3 Peltigera 11 Bacidia 3 Physcia 9 Leptogium 3 Lecanora 8 Pertusaria 3 Phaeophyscia 7 Punctelia 3 Caloplaca 5 Ramalina 3 Parmotrema 4 Stereocaulon 3 Heterodermia 4 Usnea 3 Xanthoparmelia 4 Xanthomendoza 3

Analysis of photobionts revealed that 86% are comprised of Chlorophycean algae while 8% contain cyanobacteria, and 2% utilize both green algae and cyanobacteria.

Approximately 4% of the specimens identified do not contain a symbiotic photosynthetic partner and are therefore considered not lichenized. They are however, studied by lichenologists and included in floristic surveys of lichens.

Rare and Significant Lichens

Based on the Wisconsin Natural Heritage Working List of rare species produced by the Bureau of Endangered Resources, six rare lichens were collected during this investigation. Four species, Cladonia incrassata, Peltigera elisabethae, Physcia tenella, and Punctelia caseana were all located in wet areas with Cladonia decorticata and

Peltigera extenuata found in dry – upland sites. P. extenuata is a recently segregated species from P. didactyla, a common lichen throughout Wisconsin. Since all lichens on the working list bear the state status of “special concern” no legal protection is provided.

It should be noted that all but two of these lichens were found in wet areas.

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Three other significant lichens were discovered within the Richard A. Hemp

Fishery Area which is located in eastern Portage County: Lobaria pulmonaria,

Normandina pulchella, and Leptogium rivulare. Although L. pulmonaria is not listed as rare within the state, it is of interest to find it this far south. Previously found throughout the state, in recent years it has been confined to the very northern tier of counties. This epiphytic cyanolichen is extremely sensitive to sulfur dioxide and is indicative of clean air. N. pulchella was previously reported for the state by Lay (2004) and like L. pulmonaria is considered an indicator of old growth hardwood forests in Maine (Selva

1994).

Leptogium rivulare is also an epiphytic cyanolichen and is a new report for the state (Bennett & Bartkowiak 2013). This foliose Leptogium is rarely collected because of its unusual habitat. It has only been found on the boles of trees that are periodically inundated by fresh water. Sierk (1964) mentions it only occurring on boles at pond edges, but in this case it was found on boles along a small river. Studies in Canada (COSEWIC

2004) give evidence that upstream geochemistry is important, as it is only found where water bodies are downstream of calcareous parent materials. In the WI site, the upstream geology is composed of glacial till including dolomitic pebbles (Clayton 1986).

At this site L. rivulare was found on the boles of Fraxinus nigra Marshall and

Tilia americana L. (Figure 6) it is also known to grow on Alnus, Cephalanthus, Acer,

Quercus, Thuja, Cornus, Fraxinus, Salix and Populus (Sierk 1964; Environment Canada

2012). Absent any other data on host specificity it appears the species prefers hardwoods. There appear to be a few records of it occurring on stone (Environment

Canada 2012).

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Figure 6. Habitat of L. rivulare on the Tomorrow River in Portage County.

There are only 11 records for this species in CNALH, three from the United

States, including two 19th Century specimens from Vermont, a 1974 collection from

Oregon of dubious identification, and a mis-identified 1978 collection from Georgia. The remaining North American collections include six from eastern Canada and one collection from Mexico. Wong and Brodo (1992) considered it very rare in Ontario. The remaining three collections are from Scandinavia. The species is easy to identify not only because of the unusual habitat, but also because it is the only North American species with four spored asci and no isidia. The spores are sub-muriform, with three septa vertically and one or two septa longitudinally, approximately 17-18 x 10 μ.

Cyanolichens tend to occupy relatively lower light and higher moisture habitats than lichens with green algal photobionts. As a result they are more likely to be found in

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old-growth forests, mountaintops and alpine regions, and calcareous rocks and soils

(Hinds and Hinds 2007). They are also known to fix nitrogen in these habitats and are sensitive to air pollution. The Midwestern United States currently does not contain large amounts of these habitats, and consequently cyanolichens are not abundant. Therefore sites that contain them deserve special protection to prevent their extinction. This site is managed as a fishery, which includes removal of streamside trees. This would eliminate the habitat for this species.

Wisconsin has very little old growth forests left, but calcareous habitats are abundant in several parts of the state. Since there are no alpine areas; this leaves the calcareous habitats as the most important for cyanolichens. Some of these areas include steep dolomitic bluffs, one alvar, and limestone shorelines of lakes. From this study we can add alkaline water bodies draining calcareous parent materials. These are currently not identified as a specific habitat or community type by the WI Department of Natural

Resources, although they do have categories for fast and slow, soft cold and warm streams (WI Natural Heritage Inventory). These should be particularly surveyed for cyanolichens.

Threats to L. rivulare have been reviewed by Environment Canada (2012) and include host mortality from the Emerald ash borer (Agrilus planipennis) and Dutch elm disease (Ophiostoma novo-ulmi) to the host, dusky slug, black algae, alterations of the water regime and chemistry, habitat loss, tree harvesting, water and air pollution, forest fires, ice scarring, and interspecific competition. Environment Canada already has a

Recovery Team in place to address these threats.

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Heterodermia obscurata (Nyl.) is not as rare as L. rivulare, nevertheless it is a new state record to report here. According to CNALH, distribution in North America is mainly southern and eastern with solitary collections from both Minnesota and Michigan and two from Ontario Canada. H. obscurata is a foliose lichen often found on the boles of deciduous trees and occasionally moss covered rocks in humid sites. This lichen was collected at Spring Lake in southeastern Portage County, in the Lanark Township. This species is distinguished from other Heterodermia by its ecorticate lower surface with yellow-brown patches that contain an unknown pigment and predominantly black projecting rhizines.

Peltigera extenuata is also reported here as a new occurrence in the state. It is a relatively new segregate from P. didactyla which is a common lichen of barren and sandy soil throughout the state. In 2003, Goffinet et al. fully described it as a separate species using molecular and morphological studies. The most distinguishing physical feature between the two species is the rhizines; P. extenuata possesses flocculent rhizines throughout the entire lower surface whereas P. didactyla has mostly simple rhizines and they are absent from the margins. This species was found growing directly on the soil in the Town of Hull.

In addition to the state records and rare lichens, 115 new county records are reported for Portage County with this study. This large number of county records is most likely due to the fact that the county has never before been the focus of a lichen study.

Although other lichenologists including John Thomson and William Culberson previously collected in the county, no one has thoroughly searched the area for microhabitats in an attempt to compile a thorough list.

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Culberson Site/Curtis Stand 3044 Revisited

The only study site with historical data was the Culberson site, located in northeastern Portage County in the New Hope Township. William Culberson studied corticolous lichens at this location which was also used by John Curtis in the Vegetation of Wisconsin (1959). The site was relocated by referring to the Curtis files (Stand

Number 3044) in the Plant Ecology Lab (PEL) at the University of Wisconsin – Madison.

Culberson’s data were collected during the summers of 1952 and 1953 and included only lichens found on the boles of 40 trees at breast height (4.5 feet, dbh) using a random pairs sampling method.

The property continues to belong to the Cieslewicz family as it was during the original survey. The parcel of land studied was originally entered into the Forest Crop

Law in 1950 as it was 33 acres of mature red and white pine on land that was not suitable for agriculture. The location is described as a flat topped hill with a south facing slope grading down to a lake (Figure 7).

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Figure 7. Google Earth image of Cieslewicz property. Circle indicates approximate location of Culberson study site (Curtis Stand #3044). Triangles identify two homes presently on the property.

The first recorded timber harvest following Culberson’s work occurred on the property in 1964 when 13,520 board feet were removed during a very selective cut for personal use, most likely to build a barn or a residence. A thinning of the stand occurred three years later in 1967 when 25 – 30% of the canopy was removed. The property was removed from Managed Forest Lands in 1998 (Lyle Eiden DNR Forester, personal communication). These activities would have certainly had an impact on the lichen communities.

With the property owners’ permission, the site was revisited during the summer of

2012. No attempt was made at re-surveying the location using the random pairs sampling method as Culberson had, since that was not the intention of this study.

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William Culberson identified 17 species of lichen on the boles at breast height of

40 red and white pines (Pinus resinosa and P. strobus). The survey from 2012 resulted in the identification of 34 species and 2 unidentified crustose specimens on a variety of substrates; including coniferous and deciduous tree species, granite glacial erratics, and cut stumps at various degrees of decomposition. Logging roads were evident on the property along with a small population of garlic mustard (Alliaria petiolata). Two private residences have been constructed on the property. The adjacent lands, previously used in agriculture are now red pine (Pinus resinosa) plantations. A complete list of species from this location appears in Table 3.

Twice the number of lichen species was identified in 2012 as in 1952-53. Since the sampling method in the earlier survey was limited to the boles of 40 trees at breast height, this would seem logical. However, 11 of the species identified in the previous survey were not present in 2012. Likewise, 30 new species were identified during the

2012 survey. Parmelia saxatilis reported by Culberson is suspect since it is most often a saxicolous species. P. squarrosa, a corticolous species is easily mistaken for P. saxatilis.

Inspection of the rhizines would separate the two species since P. squarrosa would have squarrosely branched rhizines and P. saxatilis with simple or furcate rhizines.

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Table 3. Lichen species identified for Culberson site, X=present. 1952- 1952- Species 1953 2012 1953 2012 Acarospora fuscata x Lepraria neglecta x Amandinea polyspora x Melanelixia subaurifera x Arthonia caesia x Myelochroa aurulenta x Aspicilia cinerea x Myelochroa galbina x Caloplaca cerina x Parmelia saxatilis x Caloplaca holocarpa x Parmelia sulcata x x Candelaria concolor x Pertusaria amara x Phaeocalicium Candelaria fibrosa x polyporaeum x Cladonia macilenta x Phaeophyscia ciliata x Cladonia coniocraea x Phaeophyscia rubropulchra x Cladonia cristatella x Physcia aipolia x Cladonia didyma x Physcia ascendens x Cladonia incrassata x Physcia millegrana x Evernia mesomorpha x x Physconia detersa x Flavoparmelia caperata x x Platismatia tuckermanii x Graphis scripta x Punctelia bollianna x Hypogymnia physodes x Punctelia rudecta x x Lecanora allophana x x Ramalina americana x Unknown Scoliciosporum crustose species 2x chlorococcum x Lecanora strobilina x Trapeliopsis flexuosa x Lecanora symmicta x Tuckermannopsis ciliaris x Lecanora thysanophora x Usnea sp. x x Lecanora varia x Xanthomendoza hasseana x Total 17 34

Among the 11 previously identified species that were absent from the 2012 survey are sensitive lichen species that may have been negatively impacted by the change in light, humidity, and air quality in the more than 60 years between samplings. The most sensitive lichen on the list is Usnea, followed by Ramalina americana. Although Usnea species were present in both samplings the only specimen found in the 2012 survey was a very small, less than 2 cm single thallus at the base of a decomposing cut stump.

William Culberson did not report any Ramalina species in his report; this may have to do

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with the light levels in the stand at that time. Very little light may have been available on the boles of the trees at breast height in a mature stand.

Lichens Previously Reported; Not Collected 2010 – 2012

Twenty-nine lichens were previously reported for Portage County that were not collected with this study. A list of those species appears in Table 4. Several of these lichens are considered rare throughout the state and may no longer be present. Usnea species in general are sensitive to air quality and it would be understandable that they are no longer found in the county. According to Thomson (2003), the most recent collection of U. angulata in the WIS herbarium is 1927. Nephroma helveticum has been collected in Portage County as recently as 1970, so with continued intense sampling this lichen quite possibly may be found again. Ramalina species are also sensitive to air quality; the only collection of R. dilacerata in the county was from 1945. Bryoria trichodes was last collected in central Wisconsin by L. Cheney at the end of the 19th Century, it is currently found in the northern counties of the state. It is likely that several of these species, especially the crustose species on the list, might be found within Portage County with continued rigorous sampling.

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Table 4. Lichens previously reported for Portage County. Amandinea dakotensis Lepraria neglecta Bryoria trichodes Melanelia olivacea Caloplaca ferruginea Nephroma helveticum Cetrelia chicitae Pertusaria macounii Cladonia cylindrica Physcia halei Cladonia humilis Porpidia macrocarpa Cladonia multiformis Punctelia hypoleucites Cladonia petrophila Ramalina dilacerata Cladonia ramulosa Rinodina pachysperma Collema nigrescens Tuckermannopsis ciliaris Fuscidea recensa var. recensa Usnea angulata Lecanora albellula Usnea mutabilis Lecanora expallens Usnea strigosa Lecanora hagenii Xanthoparmelia angustiphylla Lecanora perplexa

It is fortunate that these records are available, as they serve as baseline data for the county, one in which this study is able to build upon. It is important to note that these sensitive lichens were once present, indicating a change in air quality or forest structure. Agricultural fields have replaced many natural communities, paper mills and power plants obstruct the rivers and impact air quality, and residential neighborhoods have been developed in a variety of habitats. So many changes have occurred across the landscape in central Wisconsin; it is not surprising that many of these sensitive lichens have disappeared. Imagine paddling down the Wisconsin River before the dams obstructed the flow, when the river was wild and wove beneath conifers draped in Usnea and Bryoria species.

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COLLECTION SITES

Localities are listed in numeric then alphabetical order, followed by collection numbers of specimens.

1. 100th Street, Town of Grant. 44.322111°N, -89.687944°W. Alongside road in dry

sandy soil with Pinus banksiana in the southwest corner of the county. 570.

2. 5433 Claret Drive, Town of Hull. 44.571012°N, -89.518899°W. Field with sandy soil

in southwest portion of property. 157, 158, 163B, 432, 504.

3. 90th Street, Town of Grant. 44.301°N, -89.708083°W. Alongside road in dry sandy

soil with Pinus banksiana in the southwest corner of the county. 571-576.

4. Becker Lake. 44.60385°N, -89.4419°W. Bog lake surrounded by forest and Alder

thicket. 181-183.

5. Buena Vista Quarry Prairie SNA. 44.31942°N, -89.6286°W. Dry mesic prairie with

an oak knoll rise. 394 – 403, 410 – 412.

6. Carrie Frost Park. 44.29949°N, -89.45798°W. Managed red pine stand and school

forest. Surrounded by extensive agriculture. 761, 762.

7. Cemetery corner of CR-K & D. 44.41657°N, -89.36568°W. Collections from

headstones and sandy soil. 726, 727.

8. Cieslewicz Cedars. 44.57708°N, -89.31907°W. Cedar swamp near headwaters of

Poncho Creek. 747 – 753.

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9. Clementson Road. 44.52409°N, -89.33906°W. Flood plain area and along banks of

the Tomorrow River, within the Richard Hemp State Fishery Area, south of

confluence with Poncho Creek. 754 – 757, 763 – 766, 778, 780.

10. CR-X, west of I-39. 44.58737°N, -89.62019°W. Fraxinus sp. next to road. 44, 45.

11. Culberson site, CR-OO. 44.58944°N, -89.31006°W. Northern mesic hardwood stand.

Previously studied by William Culberson. 697 – 719.

12. Damask property. 44.57903°N, -89.3118°W. Upland forested area surrounding a

pothole lake. 301 - 328

13. Emmons Creek Barrens/Ice Age Trail. 44.304916°N, -89.234388°W. Oak savanah.

562 – 568.

14. Faraway Loop Ice Age Tail. 44.3012°N, -89.2429°W. Oak savannah uplands,

wetlands and banks of Emmons Creek. 249 – 272, 382 – 393.

15. Flume Creek State Fishery Area. 44.58542°N, -89.2954°W. Lowlands surrounding

Flume Creek. 234 – 237.

16. Fountain Lake. 44.30923°N, -89.2648°W. Hard water drainage lake, and headwaters

of Emmons Creek. 153 –156,420, 436, 783, 784.

17. Galecke Park. 44.463963°N, -89.604007°W. Wisconsin River frontage with heavy

use by water sport enthusiasts. 586.

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18. Green Circle Trail - Moses Creek. 44.562465°N, -89.539192°W. 3 mile section of

trail traversing uplands and lowlands with foot bridges through swampy areas and an

ephemeral pond. 1, 2, 29-41, 720.

19. Green Circle Trail - Plover River Trail. 44.55827°N, -89.522°W. 4 mile section of

trail traversing dry savannahs, mesic uplands, and lowlands including the Plover

River. 102-128, 130-137, 216, 421-427, 433, 455, 503, 604-606, 694-696, 728.

20. Green Circle Trail - Whiting Park. 44.49776°N, -89.53845°W. 1.5 mile section of

trail through managed Pinus resinosa stands and dry savannahs. 611-624.

21. Grover Wood Lot. 44.56205°N, -89.7946°W. Once an extremely high quality sugar

maple stand, harvested in the 1980’s. 159-162.

22. Hauer Farm, 1420 Sand Hill Lane, Town of Linwood. 44.516423°N, -89.689043°W.

Farmstead with wooded areas, fenced croplands and outbuildings. 375-381.

23. Haymeadow Drive. 44.64895°N, -89.5503°W. Dewey Marsh area. Upland area

surrounding small depression. 129, 721 – 723.

24. Iverson Park. 44.52248°N, -89.5388°W. City of Stevens Point park, includes

wetlands along the Plover River. 240-248.

25. Jacqueline Lake. 44.675776°N, -89.423354°W. Various plant communities

surrounding the lake, including conifer bogs with Picea mariana. 226-232, 284-300,

354-373, 505-509.

26. Jordan Park South . 44.57478°N, -89.5027°W. Lowlands and granite glacial erratics

along the Plover River. 85-101, 456, 457.

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27. Lions Lake. 44.646868°N, -89.285792°W. Uplands, lowlands surrounding lake. 465

& 468.

28. Little Bear Hemlocks SNA. 44.60643°N, -89.7196°W. Remnant hemlock stand. 215

29. Little Plover River, east of I-39. 44.47417°N, -89.5169°W. Uplands of mixed conifers

and hardwoods, and lowlands along the river. 273-282.

30. Little Wolf River State Fishery Area (end of Wigwam Road). 44.675194°N, -

89.233861°W. Lowlands with abundant Thuja occidentalis and granite boulders.

591-601.

31. Lone Rock. 44.24998°N, -89.5907°W. Steep-sided Cambrian sandstone outcropping.

217-225, 729-733A.

32. Mead Conifer Bogs SNA. 44.684842°N, -89.841642°W. Lowland area surrounding

the conifer bogs. 459-461.

33. Mill Road, Bancroft. 44.302283°N, -89.521766°W. Field at end of road with

deteriorating fencepost and Quercus rubra. 349-352.

34. Mosquito Bluff. 44.276611°N, -89.510305°W. Steep-sided Cambrian sandstone

outcropping. 577-585.

35. New Hope Pines SNA. 44.574333°N, -89.265333°W. Northern dry-mesic forest;

indicative of presettlement pineries with 2 kettle lakes. 587-590.

36. Northstar Road. 44.619137°N, 89.45766°W. Very large granite boulders alongside

road. 184&185.

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37. Plover River Wayside. 44.62763°N, -89.4841°W. Highway right of way, providing

access to the river; lowlands. 170-180.

38. Poncho Creek. 44.5696°N, -89.3258°W. Lowland forest (swamp conifer and

bottomland hardwoods), and mixed hardwood forest with abundant springs and seeps.

Area includes a trout stream corridor. 191-214, 331-348, 526-528.

39. Poncho Creek Hemlocks. 44.553°N, -89.329527°W. Lowland forest with abundant

hemlock and granite glacial erratic boulders. 487-498, 552-556.

40. Rosholt Swamp. 44.62767°N, -89.3639°W. Wet area with lowland conifers and

granite glacial erratics. 767-777.

41. Schmeeckle Reserve. 44.536408°N, -89.569434°W. More than 100 hectare nature

reserve with a mosaic of habitats on the UWSP campus. 472, 473, 510-516.

42. Severson Lake - Ice Age Trail. 44.54357°N, -89.23486°W. Mixed pine and hardwood

forest surrounding the lake. 653, 658-668.

43. Spring Lake. 44.392151°N, -89.339597°W. Sensitive wetlands with lowland

hardwoods and frequent springs and seeps. 438-454, 474-484, 687-692.

44. St. Peter's Cemetery. 44.53733°N, -89.59991°W. Collections from various

headstones. 607-610.

45. Stanley Street. 44.527315°N, -89.569759°W. Ash tree in boulevard on UWSP

campus. 409.

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46. Stedman Park. 44.38347°N, -89.2527°W. Old highway right-of-way with upland

woods and granite boulders along the Tomorrow River. 138-152.

47. Steinhaugen Property. 44.63789°N, -89.4668°W. 113 hectare property includes

uplands and lowlands in a managed forest habitat. 625-638, 669-686.

48. Sunset Drive, west of CR-X. 44.60527°N, -89.61866°W. West facing slope of dry

sandy soil along road. 733B-745.

49. The Nature Conservancy Park. 44.561617°N, -89.509545°W. Lowland along the

Plover River. 59-67.

50. Tomorrow River Wayside . 44.51414°N, -89.33411°W. Fishing access to the

Tomorrow River (abandoned road). 758-760.

51. Townline Road, Town of Grant. 44.278191°N, -89.606352°W. Alongside road in dry

sandy soil with Pinus banksiana in the southwest corner of the county. 569.

52. Tree Lake. 44.669666°N, -89.276055°W. Boat landing with lowlands around lake.

466, 469-471.

53. Twin Lakes Road. 44.62598°N, -89.3997°W. Hardwoods alongside road. 186-190.

54. Upper Little Wolf River SNA. 44.678861°N, -89.259222°W. Uplands and granite

boulders alongside and in the river. 529-551, 559-561, 602, 603.

55. West River Drive boat landing. 44.5063°N, -89.5941°W. Boat landing access to

Wisconsin River. 164.

45

56. West River Park. 44.50115°N, -89.5967°W. Wooded parcel across the road from the

Wisconsin River. 166-169.

57. Whiting Triangle. 44.48848°N, -89.56236°W. Marsh next to Wisconsin River. 486

58. Wolf Lake. 44.32548°N, -89.34817°W. Hard-water seepage lake with fluctuating

water tables and upland forests. 654-657.

46

CATALOGUE OF SPECIES

At the onset of this study, 67 species in 33 genera were documented for Portage County. With the conclusion of this study, the catalogue now includes 208 species in 79 genera.

The following list of lichens is according to current taxonomy and nomenclature follows Esslinger (2012). Genera and species are arranged alphabetically. Collection numbers belong to the author, followed by collection site(s) in bold. Since not all lichen taxa encountered were collected at each site, the number of collections per species does not represent abundance. If not collected by the author, the publishing source is provided and indicated with #; no attempt has been made to verify these specimens. If not previously published for the county, the collector’s name is provided in italics and the identity has been confirmed. Taxa that are new reports for Portage County are preceded by an asterisk (*); new records for the state of Wisconsin include a double asterisk (**).

* Acarospora fuscata (Schrader) Arnold, 139, 267, 403, 626A. 5, 14, 46, 47.

* Acrocordia megalospora (Fink) R. C. Harris. 450, 463, 632. 43, 47, 49.

# Amandinea dakotensis (H. Magn.) P. May & Sheard. (Thomson 2003).

* Amandinea polyspora (Willey) E. Lay & P. May. 624, 714, 701B. 11, 20.

Amandinea punctata (Hoffm.) Coppins & Scheid. 380. 22.

* Anaptychia palumulata (Michaux) Vainio. 346. 38.

Arthonia caesia (Flotow) Körber. 504C, 701A. 2, 11.

* Aspicilia caesiocinerea (Nyl. ex Malbr.) Arnold. 535, 538. 54

* Aspicilia cinerea (L.) Körber. 147, 217, 457, 512, 581, 710. 11, 26, 31, 34, 41, 46. Not previously reported, Foote 1962.

* Aspicilia laevata (Ach.) Arnold. 729A. 31

47

* Bacidia polychroa (Th. Fr.) Körber. 89. 26

* Bacidia schweinitzii (Fr. ex E. Michener) A. Schneider. 337, 553, 593, 594, 598, 601, 748. 8, 30, 38, 39

* Bacidia suffusa (Fr.) A. Schneider. 600. 30

# Bryoria trichodes (Michaux) Brodo & D. Hawksw. (Thomson 2003).

* Buellia erubescens Arnold. 302, 363. 12, 25

* Buellia schaereri De Not. 115B. 19

* Caloplaca arenaria (Pers.) Müll. Arg. 394A, 395. 5

* Caloplaca cerina (Ehrh. ex Hedwig) Th. Fr. 697B, 699A. 11

# Caloplaca ferruginea (Hudson) Th. Fr. (Thomson 2003).

Caloplaca flavorubescens (Hudson) J. R. Laundon. 474, 475, 655. 43, 58

* Caloplaca holocarpa (Hoffm. ex Ach.) A. E. Wade. 144, 697D, 699B. 11, 46

* Caloplaca microphyllina (Tuck.) Hasse. 374. 19

Candelaria concolor (Dickson) Stein. 188, 472, 609, 717. 11, 41, 44, 53

* Candelaria fibrosa (Fr.) Müll. Arg. 698, 712, 729B. 11, 31

* Candelariella efflorescens R. C. Harris & W. R. Buck. 101, 612. 20, 26

* Candelariella vitellina (Hoffm.) Müll. Arg. 219, 514. 31, 41. Not previously reported, Foote 1962.

# Cetrelia chicitae (Culb.) W. L. Culb. & C. F. Culb. (Thomson 2003).

* Chaenotheca brunneola (Ach.) Müll. Arg. 377. 22

* Cladonia arbuscula subsp. mitis (Sandst.) Ruoss. 423. 19

* Cladonia botrytis (K. Hagen) Willd. 455. 19

48

Cladonia caespiticia (Pers.) Flörke. 225, 308, 347, 440, 491, 680. 12, 31, 38, 39, 43, 47

* Cladonia cariosa (Ach.) Sprengel. 410, 571. 3, 5.

Cladonia chlorophaea (Flörke ex Sommerf.) Sprengel. 743. 48

Cladonia coniocraea (Flörke) Sprengel. 259, 327, 529, 556. 12, 14, 39, 54

* Cladonia crispata (Ach.) Flotow. 622, 694. 19, 20

Cladonia cristatella Tuck. 107, 325, 427. 12, 19

# Cladonia cylindrica (A. Evans) A. Evans. (Thomson 2003).

* Cladonia decorticata (Flörke) Sprengel. 620. 2

* Cladonia didyma (Fée) Vainio. 709. 11

Cladonia fimbriata (L.) Fr. 190, 497, 654. 39, 53, 58

Cladonia gracilis ssp. turbinata (Ach.) Ahti. 620. 20.

* Cladonia humilis (With.) J. R. Laundon. (Listed as C. conista, Thomson 2003).

* Cladonia incrassata Flörke. 486, 505, 707. 11, 25, 57

Cladonia macilenta Hoffm. 131, 354, 420, 706, 761. 6, 11, 16, 19, 25

* Cladonia metacorallifera Asahina. 370. 25

# Cladonia multiformis G. Merr. Thomson 662, WIS, listed in CNALH.

* Cladonia ochrochlora Flörke. 211, 494, 662, 752. 8, 38, 39, 42

* Cladonia parasitica (Hoffm.) Hoffm. 87, 262, 629, 663, 690. 14, 26, 42, 43, 47

Cladonia petrophila R. C. Harris. Foote 62293, WIS, listed in CNALH.

Cladonia peziziformis (With.) J. R. Laundon. 199, 722. 23, 38

49

* Cladonia phyllophora Hoffm. 565, 740, 741, 742. 13, 48

* Cladonia pyxidata (L.) Hoffm. 116, 280, 299, 546, 615, 679, 744. 19, 20, 25, 29, 47, 48, 54

Cladonia ramulosa (With.) J. R. Laundon. 138, 661. 42, 46.

* Cladonia rangiferina (L.) F. H. Wigg. 125, 265, 271, 421, 574, 734, 735. 3, 14, 19, 48

Cladonia rei (L.) F. H. Wigg. 576, 618. 3, 20

* Cladonia robbinsii A. Evans. 695. 19

* Cladonia scabriuscula (Delise) Nyl. 249, 736. 14, 48

Cladonia squamosa Hoffm. 339B, 340, 355, 361, 468, 584, 658, 678, 747, 767. 8, 25, 27, 34, 38, 40, 42, 47

* Cladonia subcariosa Nyl. 23, 126, 168, 270, 279. 14, 19, 29, 48, 56

* Cladonia symphycarpia (Flörke) Fr. 59, 572, 616, 745. 3, 19, 20, 48

* Cladonia uncialis (L.) F. H. Wigg. 21, 272, 411, 422, 562. 5, 13, 14, 19, 48

* Cladonia verticillata (Hoffm.) Ahti. 269, 281, 282, 575, 720, 733B, 738. 3, 14, 19, 29, 48

* Coenogonium luteum (Dicks.) Kalb & Lücking. 591, 773. 30, 40

* Coenogonium pineti (Ach.) Lücking & Lumbsch (Lücking, Stuart & Lumbsch 2004). 500, 749. 8, 38

# Collema nigrescens (Hudson) DC. Cheney 1898 (Thomson 1998)

Cyphelium tigillare (Ach.) Ach. 349, 379. 22, 33

* Dermatocarpon luridum (With.) J. R. Laundon. 547. 54

Diploschistes moscorum Zahlbr. 396, 424. 5, 19

50

* Diploschistes scruposus (Nyl.) Lücking, Knudsen & Fryday. 397. 5

* Eopyrenula intermedia Coppins. 453, 635. 43, 47

Evernia mesomorpha Nyl. 1, 27, 34, 35, 114, 214, 226, 290, 356, 433. 18, 19, 25, 38

* Flavoparmelia baltimorensis (Gyelnik & Fóriss) Hale. 286, 583. 25, 34

Flavoparmelia caperata (L.) Hale. 33, 37, 38, 39, 40, 41, 45, 96. 10, 18,

19, 26

Flavopunctelia flaventior (Stirton) Hale. 568. 13

* Flavopunctelia soredica (Nyl.) Hale. 175, 108. 37

* Fuscidea recensa var. recensa (Arnold) Fryday. Not previously reported, Foote 1962.

Graphis scripta (L.) Ach. 93, 339A, 483, 498, 552, 705A, 777. 11, 26, 38, 39, 40, 43

* Heterodermia galactophylla (Tuck.) Culb. 332, 336. 38

* Heterodermia hypoleuca (Ach.) Trevisan. 201, 445, 448, 592, 628. 30, 38, 43, 47

** Heterodermia obscurata (Nyl.) Trevisan. 443. 43

* Heterodermia speciosa (Wulfen) Trevisan. 130, 306, 313, 426, 479, 603, 770. 12, 19, 40, 43, 54

* Hypocenomyce scalaris (Ach.) M. Choisy. 129, 350, 674, 675. 23, 33, 47

Hypogymnia physodes (L.) Nyl. 36, 102, 111, 119, 208, 277, 287, 365, 367, 371, 672. 18, 19, 25, 29, 38, 47

* Julella fallaciosa R. C. Harris. 528, 757, 766. 9, 38

* Lecanora albella var. rubescens (Imshaug & Brodo) Lumbsch. 599. 30

# Lecanora albellula Nyl. (Thomson 2003).

Lecanora allophana Nyl. 293, 294A, 366, 715. 11, 25

51

* Lecanora caesiorubella Ach. 110, 166, 292, 360, 507, 673. 19, 25, 47, 56

# Lecanora expallens Ach. (Bennett 2006b)

# Lecanora hagenii (Ach.) Ach. Thomson 1998 – Cheney collection 1894.

* Lecanora muralis (Schreber) Rabenh. 607. 44

# Lecanora perplexa Brodo. (Thomson 2003).

Lecanora strobilina (Sprengel) Kieffer. 60, 381, 702. 11, 19, 22

* Lecanora symmicta (Ach.) Ach. 321, 563, 700. 11, 12, 13

* Lecanora thysanophora Harris. 90, 193, 335A, 705B. 11, 26, 38

Lecanora varia (Hoffm.) Ach. 551. 54

* Lepraria incana (L.) Ach. 160, 215, 503. 19, 21, 28

* Lepraria lobificans Nyl. 92, 244, 342. 24, 26, 38

# Lepraria neglecta (Nyl.) Erichsen. Culberson, 1952-3, unpublished, specimen not located.

Leptogium cyanescens (Rabenh.) Körber. 145, 204, 495, 682, 683. 38, 39, 46, 47

* Leptogium milligranum Sierk. 197, 343, 484. 38, 43

** Leptogium rivulare (Ach.) Mont. 763, 780. 9

* Lobaria pulmonaria (L.) Hoffm. 554. 39

# Melanelia olivacea (L.) O. Blanco et al. (Thomson 2003).

Melanelia septentrionalis (Lynge) O. Blanco et al. 509, 573. 3, 25

Melanelixia subaurifera (Nyl.) O. Blanco et al. 2, 61, 103, 171, 174, 256, 296, 311. 12, 14, 18, 19, 25, 37, 49

* Mycocalicium subtile (Pers.) Szatala 143, 253, 587. 14, 35, 46

52

Myelochroa aurulenta (Tuck.) Elix & Hale. 253, 323, 425, 589, 637. 12, 15, 19, 35, 47

Myelochroa galbina (Ach.) Elix & Hale. 62, 229, 316, 493. 12, 25, 39, 49

# Nephroma helveticum Ach. (Thomson 2003).

* Normandina pulchella (Borrer) Nyl. 778. 9

* Ochrolechia arborea (Kreyer) Almb. 240. 24

* Opegrapha varia Pers. 567. 13

Parmelia saxatilis (L.) Ach. 94. 26

Parmelia sulcata Taylor. 118, 122B, 173, 181, 291, 412, 459, 460, 685, 760. 4, 5, 19, 25, 32, 37, 47, 50

* Parmeliopsis ambigua (Wulfen) Nyl. 251. 14

* Parmeliopsis hyperopta (Ach.) Arnold. 449, 478. 43

* Parmotrema crinitum (Ach.) M. Choisy. 196, 345, 384, 444, 633, 775. 14, 38, 40, 43, 47

* Parmotrema hypotropum (Nyl.) Hale. 566, 668. 13, 42

* Parmotrema margaritatum (Hue) Hale. 774, 781. 9, 40

* Parmotrema reticulatum (Taylor) M. Choisy. 353, 728. 19

Peltigera canina (L.) Willd. 542, 543. 54

Peltigera didactyla (With.) J. R. Laundon. 636, 726. 7, 47

* Peltigera elisabethae Gyelnik. 492, 602, 721. 23, 39, 54

* Peltigera evansiana Gyelnik. 200, 333, 439, 688. 38, 43

** Peltigera extenuata (Vainio) Lojka. 121. 19

53

* Peltigera horizontalis (Hudson) Baumg. 545. 54

* Peltigera lepidophora (Nyl. ex Vainio) Bitter. 146, 212, 783. 16, 38, 46

* Peltigera neckeri Hepp ex Müll. Arg. 338, 437, 544, 750. 8, 38, 54

* Peltigera polydactylon (Necker) Hoffm. 334, 348, 595, 723. 23, 30, 38

* Peltigera praetextata (Flörke ex Sommerf.) Zopf. 120, 136, 137, 155, 156, 158.

2, 16, 19 Peltigera rufescens (Weiss) Humb. 16, 222, 465, 596, 660, 687, 768, 769. 19, 27, 30, 31, 40, 42, 43

* Pertusaria amara (Ach.) Nyl. 447B, 716A. 11, 43

* Pertusaria macounii (I. M. Lamb) Dibben. Not previously reported, Thomson 1972.

Pertusaria velata (Turner) Nyl. 202, 341, 344, 447A, 631. 38, 43, 47

* Pertusaria xanthodes Müll. Arg. 656. 58

* Phaeocalicium curtisii (Tuck.) Tibell. 746. 38

Phaeocalicium polyporaeum (Nyl.) Tibell. 560, 719. 11, 54

* Phaeophyscia adiastola (Essl.) Essl. 549. 54

Phaeophyscia ciliate (Hoffm.) Moberg. 657, 697A. 11, 58

* Phaeophyscia hirtella Essl. 205, 331, 454. 38, 43

* Phaeophyscia pusilloides (Zahlbr.) Essl. 189, 328, 506, 597, 608, 727. 7, 12, 25, 30, 44, 53

Phaeophyscia rubropulchra (Degel.) Essl. 32, 99, 135, 151, 456, 527. 18, 19, 26, 38, 46

* Phaeophyscia sciastra (Ach.) Moberg. 759. 50

54

* Phaeophyscia squarrosa Kashiwadani. 414, 442, 451, 452, 476, 481, 689, 756. 9,

38, 43 * Physcia adscendens (Fr.) H. Olivier. 179. 37

Physcia aipolia (Ehrh. ex Humb.) Fürnr. 44, 154, 167, 203, 234, 243, 383, 559, 697C, 711. 10, 11, 14, 15, 16, 24, 38, 54, 56

Physcia americana G. Merr. 324, 441, 630, 653, 753, 764. 8, 9, 12, 42, 43, 47

* Physcia caesia (Hoffm.) Fürnr. 501, 692. 38, 43

* Physcia dubia (Hoffm.) Lettau. 376, 758. 55, 50

Physcia halei J. W. Thomson. (Thomson 2003).

Physcia millegrana Degel. 30, 31, 105, 164, 250, 255, 273, 322. 12, 14, 18, 19, 29, 55

Physcia stellaris (L.) Nyl. 112, 132, 133, 263, 275, 471. 14, 19, 29, 52

* Physcia subtilis Degel. 218, 585, 731. 31, 34

* Physcia tenella (Scop.) DC. 194. 38

* Physciella chloantha (Ach.) Essl. 590. 35

Physconia detersa (Nyl.) Poelt. 88, 91, 100, 148, 150, 159, 178, 180, 195, 198, 209, 237, 246, 247, 304, 309, 310, 317, 326, 385, 466, 526, 704. 11, 12, 14, 15, 21, 24, 26, 37, 38, 46, 52

* Physconia perisidiosa (Erichsen) Moberg. 207, 533. 38, 54

Platismatia tuckermanii (Oakes) W. L. Culb. & C. F. Culb. 66. 49

* Pleopsidium flavum (Bellardi) Körber. 733A. 31

* Polysporina urceolata (Anzi) Brodo. 580, 666. 34, 42

* Porpidia albocaerulescens (Holl ex Nyl.) Henssen. 487. 39

# Porpidia macrocarpa (DC.) Hertel & A. J. Schwab. (Thomson 2003).

55

Punctelia bolliana (Müll. Arg.) Krog. 127, 128, 186, 216, 236, 257, 261, 283, 351, 352, 389. 14, 15, 19, 33, 53

* Punctelia caseana Lendemer & Hodkinson. 373. 25

* Punctelia hypoleucites (Nyl.) Krog. Thomson 16676, WIS, listed in CNALH.

Punctelia rudecta (Ach.) Krog. 95, 97, 122A, 169, 176, 192, 221, 241, 245, 260, 320, 358, 477, 550. 12, 14, 19, 24, 25, 26, 31, 37, 38, 43, 54, 56

* Pyrenula pseudobufonia (Rehm) R. C. Harris. 665. 42

* Pyxine sorediata (Ach.) Mont. 210, 446. 38, 43

Ramalina americana Hale. 63, 64, 124, 227, 228, 264, 278, 436, 461, 564, 578, 718. 11, 13, 14, 16, 19, 25, 29, 32, 34, 49

# Ramalina dilacerate (Hoffm.) Hoffm. (Thomson 2003).

* Ramalina intermedia Delise ex Nyl.) Nyl. 220. 31

* Ramalina sinensis Jatta. 182, 187, 224, 284, 305, 307, 611. 4, 12, 20, 25, 31, 53

* Rhizocarpon disporum (Nägeli ex Hepp) Müll. Arg. 511. 41

* Rhizoplaca subdiscrepans (Nyl.) R. Sant. 510, 586. 17, 41

* Rinodina oxydata (Th. Fr.) Hertel & Rambold. 399, 400. 5

# Rinodina pachysperma H. Magn. (Thomson 2003).

* Sarcogyne privigna (Ach.) A. Massal. 579. 34

* Sarea resinae (Fr.) Kuntze. 771. 40

Scoliciosporum chlorococcum (Stenh.) Vězda. 606. 19

* Stenocybe pullatula (Ach.) Stein. 29. 18

* Stereocaulon paschale (L.) Hoffm. 619. 20

56

Stereocaulon saxatile H. Magn. 784. 16

Stereocaulon tomentosum Fr. 386. 14

* Strigula submuriformis (R. C. Harris) R. C. Harris. 161, 453, 536, 627. 21, 43, 47, 54

* Trapelia coarctata (Turner ex Sm. & Sow.) M. Choisy. 402. 5

* Trapeliopsis flexuosa (Fr.) Coppins & P. James. 294B, 716B. 11, 25

* Tuckermanella fendleri (Nyl.) Essl. 106, 109. 19

Tuckermannopsis americana (Sprengel) Hale. 67, 104, 515, 570, 605. 1, 19, 41, 49

# Tuckermannopsis ciliaris (Ach.) Gyelnik. (Thomson 2003).

* Tuckermannopsis orbata (Nyl.) M. J. Lai. 65, 285, 415. 25, 49

# Usnea angulata Ach. (Thomson 2003).

* Usnea filipendula Stirton. 242. 24

Usnea hirta (L.) F. H. Wigg. 232, 390, 671. 14, 25, 47

# Usnea mutabilis Stirton. Thomson 1998 - Cheney collection 1894.

# Usnea strigosa (Ach.) Eaton. (Thomson 2003).

Usnea subfloridana Stirton. 26, 117, 230, 231, 300, 391. 14, 18, 19, 25

* Verrucaria glaucovirens Grummann. 488, 539. 39, 54

Xanthomendoza fallax (Hepp) Søchting, Kärnefelt & S. Kondr. 266, 409. 14

Xanthomendoza hasseana (Räsänen) Søchting, Kärnefelt & S. Kondr. 152, 504A, 613, 697E. 2, 11, 20, 46

Xanthomendoza ulophyllodes (Gyelnik) Søchting, Kärnefelt & S. Kondr. 85, 141, 149, 162, 170, 254, 319, 382. 12, 14, 21, 26, 37, 46

# Xanthoparmelia angustiphylla (Gyelnik) Hale. (Thomson 2003).

57

* Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale. 184, 289, 638. 25, 36, 47

Xanthoparmelia cumberlandia (Gyelnik) Hale. 298, 532. 25, 54

Xanthoparmelia plittii (Gyelnk) Hale. 223, 258, 626B. 14, 31, 47

Xanthoparmelia viriduloumbrina (Gyelnik) Lendemer. 185, 387, 392, 393. 14, 36

* Xanthoria elegans (W. A. Weber) Kalb & Hafellner. 610. 44

Xanthoria polycarpa (Hoffm.) Rieber. 113, 163B, 183. 2, 4, 19

58

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