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4.5: Macrofungi - Biology

4.5: Macrofungi - Biology


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Ascomycota

Learning Objectives

  • Use life history traits and morphological features to distinguish between Ascomycota and Basidiomycota.
  • Identify structures in the Ascomycota life cycle and know their ploidy.
  • Differentiate between different types of ascocarps; locate fertile surfaces within those structures.

The majority of described fungal species belong to the Phylum Ascomycota. Fungi in this group have simple septate hyphae and most produce fruiting bodies called ascocarps (Figure (PageIndex{1})) for sexual reproduction. Some genera of Ascomycota either only reproduce asexually via conidia or the sexual phases have not been discovered or described. These were referred to as the Fungi Imperfecti or Deuteromycota. When ascomycetes reproduce sexually, they produce haploid ascospores (usually 8) within a sac-like structure called an ascus (Figure (PageIndex{2})).

The Ascomycota include some ectomycorrhizal fungi (e.g. truffles and morels), fungi that are used as food, others that are common causes of food spoilage (bread molds and plant pathogens), and still others that are human pathogens. Some notable examples of ascomycetes include:

  • Saccharomyces cerevisiae one of the budding yeasts. It ferments sugar to ethanol and carbon dioxide and thus is used to make alcoholic beverages like beer and wine, to make ethanol for industrial use and in baking (it is often called baker's yeast). Here, it is the carbon dioxide that is wanted (to make bread and cakes "rise" and have a spongy texture). Yeast is also used in the commercial production of some vitamins and in the production - using recombinant DNA technology - of some human therapeutic proteins.
  • Neurospora crassa, another favorite "model" organism in the laboratory.
  • The fungal partner in most lichens is an ascomycete.
  • Powdery mildews that attack ornamental plants.
  • The chestnut blight, which in a few decades killed almost all of the mature American chestnut trees in the Appalachians of North America.
  • The Dutch elm disease, which has killed many of the American elms in the United States.
  • Pneumocystis jirovecii, which is a major cause of illness in immunosuppressed people, e.g., patients with AIDS.
  • The truffle and the morel, both highly-prized food delicacies. Truffles establish a symbiotic relationship with the roots of such trees as oaks.

Ascomycete Life Cycle

Asexual reproduction is frequent and involves the production of conidiophores that release haploid conidia. Sexual reproduction starts with the development of special hyphae from either one of two types of mating strains. One strain produces an antheridium and a strain of a complementary mating type develops an ascogonium. At fertilization, the antheridium and the ascogonium combine in plasmogamy without nuclear fusion. Dikaryotic ascogenous hyphae arise, in which pairs of nuclei migrate: one from each parent strain. In each ascus mother cell, two nuclei fuse (karyogamy). During sexual reproduction, thousands of asci may fill a fruiting body called the ascocarp. The diploid nucleus gives rise to four haploid nuclei by meiosis. In most ascomycetes, this is followed by a round of mitosis, producing 8 ascospores. The ascospores are then released, germinate, and form haploid hyphae that are disseminated in the environment and start new mycelia (Figure (PageIndex{4})).

Which of the following statements is true?

  1. A dikaryotic ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
  2. A diploid ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
  3. A haploid zygote that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
  4. A dikaryotic ascus that forms in the ascocarp undergoes plasmogamy, meiosis, and mitosis to form eight ascospores.

Types of Ascocarps

Apothecium

Apothecia are cup-shaped with the asci fully exposed, lining the interior of the cup. Normally, these asci are microscopic. However, in the fungus Ascobolus, the large asci with dark ascospores can be seen with the naked eye or a handlens (Figure (PageIndex{5})). The typical cup shape can be inverted and take on strange morphologies (see Figure (PageIndex{6})).

Perithecium

A perithecium is a flask-shaped fruiting structure (Figure (PageIndex{7})), often microscopic and embedded within either the substrate it is fruiting in or a fungal structure called a stroma (Figure (PageIndex{8})). The asci are almost entirely closed off from the external environment, excepting a small hole in the top of the perithecium called an ostiole. Many plant parasites, such as the causal agents of Dutch elm disease, American chestnut blight, and ergot, are perithecial ascomycetes.

Cleistothecium

A cleistothecium is a fully-enclosed fruiting structure. These typically have bag-like asci (Figure (PageIndex{9})). Some (chasmothecia) split open to release their spores. This type of fruiting body can be found in the powdery mildews, a group of plant pathogenic fungi in the order Erysiphales (Figure (PageIndex{10})).

Lichenized Fungi

Learning Objectives

  • Explain the lichen symbiosis.
  • Identify structures in the lichen thallus.

Lichens are fungi that live in a symbiotic association with a green alga or cyanobacterium (the "photobiont"), or both. The primary fungal partner (the "mycobiont") in most lichens (98% of them) is an ascomycete. Basidiomycetes make up the remainder. The relationship is often characterized as mutualistic; that is, both partners benefit. However, evidence (see "The British Soldier" below) suggests that while the fungus is dependent on its autotrophic partner, the photobiont is often perfectly content to live alone. Recently many lichens have been found to harbor a second fungal partner, a basidiomycete yeast. Its function remains to be discovered, though the presence of the basidiomycete can change the outward appearance of the lichen (e.g. color).

The British Solder

The below image is of the colorful lichen called British soldier. The fungus is Cladonia cristatella, an ascomycete. Its name is the name given to the lichen. The photobiont is Trebouxia erici, a green alga. It is found in many other lichens as well, and also can be found growing independently. The algal cells eventually are killed by the fungus, but are continuously replaced by new ones. So, the relationship in this lichen is one of controlled parasitism rather than mutualism.

The red cap produces the spores of the fungus, but these alone cannot form new lichens. Other structures (e.g., soredia), containing both partners, are needed to disperse the lichen to new locations. Some lichens release only fungal spores. These mycobionts depend for their continued survival on finding an acceptable photobiont released from other lichens. Phylogenetic trees, based on both ribosomal RNA genes and many protein-coding genes, as well as fossils indicate that lichens have been present on the earth for at least 600 million years.

Today about 14,000 species of fungi are known to form lichens. Lichens display a range of colors and textures and can survive in the most unusual and hostile habitats, though they are extremely sensitive to air pollution.. They cover rocks, gravestones, tree bark, and the ground in the tundra where plant roots cannot penetrate. Lichens can survive extended periods of drought, become completely desiccated, and then rapidly become active once water is available again. In part because of this ability, as well as the pigments in some lichens that protect from UV radiation, lichens are some of the only living things to survive exposure to conditions in space.

Link to Learning

Explore the world of lichens using this site from Oregon State University.

The body of a lichen, referred to as a thallus (Figure (PageIndex{13})), is formed of hyphae wrapped around the photosynthetic partner. The photosynthetic organism provides carbon and energy in the form of carbohydrates. If cyanobacteria are involved, they fix nitrogen from the atmosphere, contributing nitrogenous compounds to the association. In return, the fungus supplies minerals and protection from dryness and excessive light by encasing the algae in its mycelium. The fungus also attaches the symbiotic organism to the substrate.

The thallus of lichens grows very slowly, expanding its diameter a few millimeters per year. Both the fungus and the alga participate in the formation of dispersal units for reproduction. Some lichens produce soredia (Figure (PageIndex{14})), clusters of algal cells surrounded by mycelia, for asexual reproduction. Soredia are dispersed by wind and water and form new lichens. To reproduce sexually, the fungus makes spores that must find a new photosynthetic partner shortly after germinating.

Basidiomycota

Learning Objectives

  • Use life history traits and morphological features to distinguish between Ascomycota and Basidiomycota
  • Identify the parts of a mushroom
  • Identify structures in the Basidiomycota life cycle and know their ploidy

The Basidiomycota (basidiomycetes) are fungi that produce haploid basidiospores (spores produced through budding) from club-shaped cells called basidia (Figure (PageIndex{13})). These are typically formed within fruiting bodies called basidiocarps. They are important as decomposers (particularly of wood), plant pathogens, mutualists, and food sources for many animals. Basidiomycetes include the group of fungi that forms mushrooms, the rusts, and the smuts. Mushrooms are basidiocarps formed from masses of interwoven hyphae growing up from the mycelium. The basidia develop on fertile surfaces of the mushroom and release their spores (typically four from each basidium) into the air.

They hyphae of basidiomycetes are septate with clamp connections where the septa form (Figure (PageIndex{14})). The septal structure is more complex than in ascomycetes. The septum is swollen around the pore (a dolipore septum) and is flanked by structures called parenthesomes (Figure (PageIndex{15})).

Fungi with the following structures can be placed in the Basidiomycota*:

*It is important to note that these features may look different or not be present at all in some groups of Basidiomycota, such as the rusts (Pucciniomycotina) and smuts (Ustilagomycotina). However, there are many physiological and genetic similarities that support grouping these organisms together in the Basidiomycota.

Basidia and Basidiospores

Both karyogamy and meiosis occur within a cell called the basidium (Figure (PageIndex{15})). Haploid basidiospores from atop projections on the basidum called sterigmata (sing. sterigma). There are generally four spores, as shown in the image below, though the number of spores produced can vary by species. For example, the mushroom you are likely most familiar with, Agaricus bisporus (though you probably know it as a crimini or button mushroom at its immature stage and portobello at maturity), only produces two spores on each basidium (bi- meaning two).

Clamp Connections

Basidiomycetes maintain their dikaryotic (n+n) state in each hyphal compartment by making structures called clamp connections (Figure (PageIndex{16})). These are not always present, but provide a helpful identification feature when they are!

Complex Septations

Complex septations are shown in Figure (PageIndex{17}).

General Mushroom Anatomy

Though only a subset of basidiocarps look this way, they are the model for how we describe "mushrooms". In mycology, this type of basidiocarp is called "agaricoid" or "agaric" because it is the general form we see in the genus Agaricus. A more complex version of the agaric mushroom is seen in the genus Amanita (Figure (PageIndex{18})).

Basidiomycete Life Cycle

The life cycle of basidiomycetes involves an extension of the dikaryotic phase (Figure (PageIndex{19})). Mycelia of different mating strains combine (plasmogamy) soon after germination and produce a secondary, dikaryotic mycelium that contains haploid nuclei of two different mating strains (a dikaryon). This is the dikaryotic stage of the basidiomycete life cycle, and it is the dominant stage. Eventually, the secondary mycelium generates a basidiocarp. The basidiocarp can vary greatly in morphology, but in the sense of a standard mushroom (Figure (PageIndex{15})), the developing basidia are produced on the surface of gills located under cap.Within the club-shaped basidium, meiosis occurs and a diploid zygote is formed (karyogamy). The zygote divides by meiosis to produce four haploid nuclei. The haploid nuclei migrate into basidiospores, which germinate and generate monokaryotic hyphae. The mycelium that results is called a primary mycelium.

Summary

Macrofungi are represented by two major groups that can form macroscopic fruiting structures. However, many lineages within these groups might only reproduce asexually, form yeasts, or form microscopic fruiting structures (such as the rusts and smuts).

Ascomycetes typically form 8 ascospores within a structure called an ascus. In most lineages, these are produced within an ascocarp, which can be an apothecium (cup), perithecium (flask), or cleistothecium (ball). Ascomycetes have hyphae with simple septations. Their life cycle involves an extended dikaryotic phase that takes place within the ascocarp, forming dikaryotic ascogenous hyphae. These hyphae will eventually form asci, where karyogamy will take place, followed shortly after by meiosis, and usually mitosis.

Basidiomycetes typically form 4 basidiospores externally on a basidium. In the Agaricomycotina, these are produced on or in a basidiocarp, what we call mushrooms, specialized for spore dispersal. The basidiomycete life cycle is almost entirely dikaryotic. Haploid spores germinate, form a monokaryon, then must fuse with another monokaryon shortly afterward. These forms a dikaryotic mycelium with dolipore septations and clamp connections. Karyogamy only takes place within the basidia, follow promptly by meisosis to produce basidiospores.

Both ascomycetes and basidiomycetes for relationships with photosynthetic partners, such as algae or cyanobacteria, to form lichens. In this mutualism, the photobiont provides sugars from photosynthesis, while the mycobiont forms a protective thallus. In the case of cyanobacteria, these may also provide nitrogen fixation.


4.5: Macrofungi - Biology

All articles published by MDPI are made immediately available worldwide under an open access license. No special permission is required to reuse all or part of the article published by MDPI, including figures and tables. For articles published under an open access Creative Common CC BY license, any part of the article may be reused without permission provided that the original article is clearly cited.

Feature Papers represent the most advanced research with significant potential for high impact in the field. Feature Papers are submitted upon individual invitation or recommendation by the scientific editors and undergo peer review prior to publication.

The Feature Paper can be either an original research article, a substantial novel research study that often involves several techniques or approaches, or a comprehensive review paper with concise and precise updates on the latest progress in the field that systematically reviews the most exciting advances in scientific literature. This type of paper provides an outlook on future directions of research or possible applications.

Editor’s Choice articles are based on recommendations by the scientific editors of MDPI journals from around the world. Editors select a small number of articles recently published in the journal that they believe will be particularly interesting to authors, or important in this field. The aim is to provide a snapshot of some of the most exciting work published in the various research areas of the journal.


Species Richness and Traditional Knowledge of Macrofungi (Mushrooms) in the Awing Forest Reserve and Communities, Northwest Region, Cameroon

Macrofungi are diverse in their uses as food and medicine and several species serve as decomposers and also form mycorrhizal associations. Awing forest reserve is diverse in plants and fungi species. However, no work has been carried out to assess the diversity and traditional knowledge of macrofungi in the area. Diversity surveys were carried out in three altitudes using transects of

m for six months in 2015. Ethnomycology studies were carried out in fifteen communities using focus group discussion, pictorial presentation, and questionnaires. The data was analyzed using descriptive statistics in Microsoft Excel 2010. Seventy-five species belonging to thirty families were identified by morphology. Thirty-six species were found only in the low altitude, 16 in the mid altitude, and 16 species in high altitude. One species was common to low and mid altitude and also low and high altitude five species were common to mid and high altitude while there was no species common to all three altitudes. The indigenes of the Awing communities commonly called mushroom “Poh” and use it mainly as food and medicine and in mythological beliefs. The most utilized species as food and medicine included Termitomyces titanicus, Laetiporus sulphureus, and Ganoderma sp.

1. Introduction

Fungi are the most diverse organisms on earth and are defined as a eukaryotic, heterotrophic which is devoid of chlorophyll and obtains its nutrients by absorption and reproduces by means of spores [1]. Large fungi are those that form large fructifications visible without the aid of the microscope and include Basidiomycota and Ascomycota with large observable spore bearing structures [2, 3]. Ecologically, macrofungi can be classified into three groups: the saprophytes, the parasites, and the symbiotic (mycorrhizal) species. Most terrestrial fungi are saprobes or mycorrhizal symbionts, but some are pathogens of plants or fungi. Macrofungi fruiting on woody substrate are usually either saprobes or plant pathogens [4, 5]. Fungi of various taxonomic groups producing conspicuous sporocarps are collectively known as macrofungi which include “gilled fungi,” “jelly fungi,” “coral fungi,” “stink fungi,” “bracket fungi,” “puffballs,” “truffles,” and “birds nest” [6]. Macrofungal diversity is an important component of the global diversity, particularly community diversity, which is an essential part of fungal diversity [7]. Mushrooms are widespread in nature and they still remain the earliest form of fungi known to mankind [8].

Only about 6.7% of the 1.5 million species of fungi estimated in the world have been described and these are mostly in temperate regions. The tropical region which has the highest fungal diversity has not been fully exploited [9]. Cameroon has a rich biodiversity but it remains poorly unexplored. Termitomyces spp. are widely distributed across the country and form an important source of income for the rural people of Baligham and Ndop plains of the Northwest Region of Cameroon as well as Mbouda in the Western part of the country [10]. Checklist of macrofungi of Mount Cameroon consisted of 177 species as reported by [11].

Wild edible mushrooms are one of the most important natural resources on which the people of many nationalities rely and play a key role in nutrition [12]. Ethnomycology investigates the indigenous knowledge of mushroom utilization and consumption patterns such as in nutrition, medicine, and other uses [13]. It also investigates the ectomycorrhizal association and ecological benefits of macrofungi (mushrooms) to the forest. In Cameroon, mushrooms are known and consumed in many households, in the country sides and in forest areas [13]. During the onset of the rainy season when mushrooms are abundant most people in the rural areas collect them from the forest for consumption and sale [10]. The current rate of bush burning, deforestation, and overexploitation of both timber and nontimber products are threatening mushroom diversity in Cameroon. The use of fungi for food and medicine goes back a long way in human history, but research and documentation of such knowledge are relatively new in Cameroon even though one hundred and seventy-seven species of mushroom were identified in the Mount Cameroon Region [11]. Based on literature available to us macrofungi diversity in the Awing forest reserve has not been studied and there is no documentation on their ethnomycological knowledge. It is therefore crucial to document the diversity and ethnomycology of macrofungi in the Awing forest reserve and communities. Hence the objective of this study was to investigate the mushroom species richness in the Awing forest reserve with the aim of producing a checklist of macrofungi for the area and also to document the traditional knowledge of mushrooms in the communities surrounding the Awing forest reserve.

2. Materials and Methods

2.1. Study Area

Awing is found in the Northwest Region of Cameroon in West-Central Africa. It is located between latitude 05°51.527′N and longitude 010°12.122′E, with an altitude of 2126 m. Awing has a surface area of about 100 km 2 . The climate is tropical with dry and rainy seasons. It has a humidity of 98% and it is a grass-field area with fertile volcanic soils. The map of the sampled area in the Awing forest reserves for diversity studies is found in Figure 1 while the map of the sampled area in the Awing communities used for ethnomycological studies is shown in Figure 2.


About the Author

Dr. Bhim Pratap Singh working as Assistant Professor in the Department of Biotechnology, Aizawl, Mizoram University, India, has more than 10 years of research experience in the field of molecular microbiology, biocontrol of plant diseases, mushroom diversity, biosynthetic potential of actinobacteria and fungi associated with medicinal plants. He has worked for last 10 years on microbial diversity and explored the microbial population associated with medicinal plants and rhizospheric soils. His group has documented the wild mushroom of from Mizoram, Northeast India. He has completed seven externally funded research projects on screening and characterization of endophytic actibacteria and fungi associated with medicinal plants and the rhizospheric soils funded by several funding agencies. Dr. Singh has organized several international and national conferences and seminars including India-UK scientific seminar jointly funded by the British Council and the Department of Science and Technology (DST), New Delhi. Dr. Singh has been awarded with several honors like young scientist award in 2014 from Scientific and education Research Society, India and Best achievement award for the contribution to the Asian PGPR society, USA. Dr. Singh is serving ad editors of several reputed peer reviewed scientific journals like Frontiers in Microbiology, Plos ONE etc., and he is an active member of several national and international professional bodies like Asian PGPR Society, Association of Microbiologists of India, MSI, ISCA etc.. His group has published more than 45 research papers in SCI journals and 16 book chapters in the books published by national and international publishers. Dr. Singh has also edited two books published from Springer and Elsevier publications. Dr. Singh has guided 4 Ph.D students and 3 M.Phil students till date.

Lallawmsanga Chhakchhuak

Mr. Lallawmsanga is pursuing Ph.D. at the Department of Biotechnology, Mizoram University. With his research experience at the Centre for advanced studies in Botany, University of Madras, India. He has been working in the field of plant science and environmental microbiology during the last eight years. He has attended several national and international conferences. Mr. Lallawmsanga has published one paper in PLOSONE international journal to explain about diversity and antimicrobial potential of wild edible mushroom from Mizoram, Northeast India. He has been actively taking part in the university outreach programmes to educate the tribal community in Mizoram, India.

Dr. Ajit Kumar Passari working as Research Associate in Department of Biotechnology, Aizawl, Mizoram University, India, has more than six years of research experience on various fields of molecular marker study and their phylogenetic analysis, Molecular biology, secondary metabolites production and plant growth promoting potential of actinobacteria associated with medicinal plant. He has especially worked on DNA fingerprinting of endophytic actinobacteria, fungi and mushroom and screening for their antimicrobial biosynthetic potential. His research team has explored number of actinobacteria, fungi and mushroom from different sources of Mizoram, India and their functional approach. Dr. Passari has been received various awards like Young Scientist Award in International Conference on “Innovative Approaches in Applied Sciences and Technologies” at Nanyang Technological University, Singapore and selected for 2 nd BRICS Young Scientist Forum in 2017 at Zhejiang University in Hangzhou, China. He is an active member of various national and international societies like Asian PGPR Society of Sustainable Agriculture, Association of Microbiologists of India and Indian Science Congress etc. Dr. Passari has published more than 25 research papers in peer reviewed national and international journals with high impact factor. He has published 08 book chapters in national and international publishers as well as also edited one book in actinobacteria: diversity and biotechnology applications published from Elsevier publications. --This text refers to the hardcover edition.

From the Back Cover

Mushrooms are fleshy fungi with a high prospective for the production of secondary metabolites including extracellular enzymes with high agricultural and biotechnological significance. Worldwide, they are well recognized as supplementary foods due to their high nutritional values and their medicinal importance, which includes their uses in exhibiting antioxidant and antimicrobial activities, immune enhancer, and to be effective for the treatment of several diseases including diabetes and few types of cancers as well. According to recent studies, extracellular enzymes produced by several white-rot fungal strains such as Phanerochaete chrysosporium, Pleurotus sajor-caju and several mushrooms have shown a high capacity to decolorize dyes that are very harmful for the environment. Moreover, wild macrofungi have the capability to synthesize nanoparticles which are more useful for the treatment of cancer, gene therapy, DNA analysis and biosensors. Wild macrofungi are extremely important model for basic biology and commercial manufacture.


4.5: Macrofungi - Biology

All articles published by MDPI are made immediately available worldwide under an open access license. No special permission is required to reuse all or part of the article published by MDPI, including figures and tables. For articles published under an open access Creative Common CC BY license, any part of the article may be reused without permission provided that the original article is clearly cited.

Feature Papers represent the most advanced research with significant potential for high impact in the field. Feature Papers are submitted upon individual invitation or recommendation by the scientific editors and undergo peer review prior to publication.

The Feature Paper can be either an original research article, a substantial novel research study that often involves several techniques or approaches, or a comprehensive review paper with concise and precise updates on the latest progress in the field that systematically reviews the most exciting advances in scientific literature. This type of paper provides an outlook on future directions of research or possible applications.

Editor’s Choice articles are based on recommendations by the scientific editors of MDPI journals from around the world. Editors select a small number of articles recently published in the journal that they believe will be particularly interesting to authors, or important in this field. The aim is to provide a snapshot of some of the most exciting work published in the various research areas of the journal.


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Global diversity and distribution of macrofungi. / Mueller, Gregory M. Schmit, John P. Leacock, Patrick R. Buyck, Bart Cifuentes, Joaquín Desjardin, Dennis E. Halling, Roy E. Hjortstam, Kurt Iturriaga, Teresa Larsson, Karl Henrik Lodge, D. Jean May, Tom W. Minter, David Rajchenberg, Mario Redhead, Scott A. Ryvarden, Leif Trappe, James M. Watling, Roy Wu, Qiuxin.


Contents

The terms "mushroom" and "toadstool" go back centuries and were never precisely defined, nor was there consensus on application. During the 15th and 16th centuries, the terms mushrom, mushrum, muscheron, mousheroms, mussheron, or musserouns were used. [2]

The term "mushroom" and its variations may have been derived from the French word mousseron in reference to moss (mousse). Delineation between edible and poisonous fungi is not clear-cut, so a "mushroom" may be edible, poisonous, or unpalatable. [3] [4] The word toadstool appeared first in 14th century England as a reference for a "stool" for toads, possibly inferring an inedible poisonous fungus. [5]

Identifying mushrooms requires a basic understanding of their macroscopic structure. Most are Basidiomycetes and gilled. Their spores, called basidiospores, are produced on the gills and fall in a fine rain of powder from under the caps as a result. At the microscopic level, the basidiospores are shot off basidia and then fall between the gills in the dead air space. As a result, for most mushrooms, if the cap is cut off and placed gill-side-down overnight, a powdery impression reflecting the shape of the gills (or pores, or spines, etc.) is formed (when the fruit body is sporulating). The color of the powdery print, called a spore print, is used to help classify mushrooms and can help to identify them. Spore print colors include white (most common), brown, black, purple-brown, pink, yellow, and creamy, but almost never blue, green, or red. [6]

While modern identification of mushrooms is quickly becoming molecular, the standard methods for identification are still used by most and have developed into a fine art harking back to medieval times and the Victorian era, combined with microscopic examination. The presence of juices upon breaking, bruising reactions, odors, tastes, shades of color, habitat, habit, and season are all considered by both amateur and professional mycologists. Tasting and smelling mushrooms carries its own hazards because of poisons and allergens. Chemical tests are also used for some genera. [7]

In general, identification to genus can often be accomplished in the field using a local mushroom guide. Identification to species, however, requires more effort one must remember that a mushroom develops from a button stage into a mature structure, and only the latter can provide certain characteristics needed for the identification of the species. However, over-mature specimens lose features and cease producing spores. Many novices have mistaken humid water marks on paper for white spore prints, or discolored paper from oozing liquids on lamella edges for colored spored prints.

Typical mushrooms are the fruit bodies of members of the order Agaricales, whose type genus is Agaricus and type species is the field mushroom, Agaricus campestris. However, in modern molecularly defined classifications, not all members of the order Agaricales produce mushroom fruit bodies, and many other gilled fungi, collectively called mushrooms, occur in other orders of the class Agaricomycetes. For example, chanterelles are in the Cantharellales, false chanterelles such as Gomphus are in the Gomphales, milk-cap mushrooms (Lactarius, Lactifluus) and russulas (Russula), as well as Lentinellus, are in the Russulales, while the tough, leathery genera Lentinus and Panus are among the Polyporales, but Neolentinus is in the Gloeophyllales, and the little pin-mushroom genus, Rickenella, along with similar genera, are in the Hymenochaetales.

Within the main body of mushrooms, in the Agaricales, are common fungi like the common fairy-ring mushroom, shiitake, enoki, oyster mushrooms, fly agarics and other Amanitas, magic mushrooms like species of Psilocybe, paddy straw mushrooms, shaggy manes, etc.

An atypical mushroom is the lobster mushroom, which is a deformed, cooked-lobster-colored parasitized fruitbody of a Russula or Lactarius, colored and deformed by the mycoparasitic Ascomycete Hypomyces lactifluorum. [8]

Other mushrooms are not gilled, so the term "mushroom" is loosely used, and giving a full account of their classifications is difficult. Some have pores underneath (and are usually called boletes), others have spines, such as the hedgehog mushroom and other tooth fungi, and so on. "Mushroom" has been used for polypores, puffballs, jelly fungi, coral fungi, bracket fungi, stinkhorns, and cup fungi. Thus, the term is more one of common application to macroscopic fungal fruiting bodies than one having precise taxonomic meaning. Approximately 14,000 species of mushrooms are described. [9]

A mushroom develops from a nodule, or pinhead, less than two millimeters in diameter, called a primordium, which is typically found on or near the surface of the substrate. It is formed within the mycelium, the mass of threadlike hyphae that make up the fungus. The primordium enlarges into a roundish structure of interwoven hyphae roughly resembling an egg, called a "button". The button has a cottony roll of mycelium, the universal veil, that surrounds the developing fruit body. As the egg expands, the universal veil ruptures and may remain as a cup, or volva, at the base of the stalk, or as warts or volval patches on the cap. Many mushrooms lack a universal veil, therefore they do not have either a volva or volval patches. Often, a second layer of tissue, the partial veil, covers the bladelike gills that bear spores. As the cap expands, the veil breaks, and remnants of the partial veil may remain as a ring, or annulus, around the middle of the stalk or as fragments hanging from the margin of the cap. The ring may be skirt-like as in some species of Amanita, collar-like as in many species of Lepiota, or merely the faint remnants of a cortina (a partial veil composed of filaments resembling a spiderweb), which is typical of the genus Cortinarius. Mushrooms lacking partial veils do not form an annulus. [10]

The stalk (also called the stipe, or stem) may be central and support the cap in the middle, or it may be off-center and/or lateral, as in species of Pleurotus and Panus. In other mushrooms, a stalk may be absent, as in the polypores that form shelf-like brackets. Puffballs lack a stalk, but may have a supporting base. Other mushrooms, such as truffles, jellies, earthstars, and bird's nests, usually do not have stalks, and a specialized mycological vocabulary exists to describe their parts.

The way the gills attach to the top of the stalk is an important feature of mushroom morphology. Mushrooms in the genera Agaricus, Amanita, Lepiota and Pluteus, among others, have free gills that do not extend to the top of the stalk. Others have decurrent gills that extend down the stalk, as in the genera Omphalotus and Pleurotus. There are a great number of variations between the extremes of free and decurrent, collectively called attached gills. Finer distinctions are often made to distinguish the types of attached gills: adnate gills, which adjoin squarely to the stalk notched gills, which are notched where they join the top of the stalk adnexed gills, which curve upward to meet the stalk, and so on. These distinctions between attached gills are sometimes difficult to interpret, since gill attachment may change as the mushroom matures, or with different environmental conditions. [11]

Microscopic features

A hymenium is a layer of microscopic spore-bearing cells that covers the surface of gills. In the nongilled mushrooms, the hymenium lines the inner surfaces of the tubes of boletes and polypores, or covers the teeth of spine fungi and the branches of corals. In the Ascomycota, spores develop within microscopic elongated, sac-like cells called asci, which typically contain eight spores in each ascus. The Discomycetes, which contain the cup, sponge, brain, and some club-like fungi, develop an exposed layer of asci, as on the inner surfaces of cup fungi or within the pits of morels. The Pyrenomycetes, tiny dark-colored fungi that live on a wide range of substrates including soil, dung, leaf litter, and decaying wood, as well as other fungi, produce minute, flask-shaped structures called perithecia, within which the asci develop. [12]

In the Basidiomycetes, usually four spores develop on the tips of thin projections called sterigmata, which extend from club-shaped cells called a basidia. The fertile portion of the Gasteromycetes, called a gleba, may become powdery as in the puffballs or slimy as in the stinkhorns. Interspersed among the asci are threadlike sterile cells called paraphyses. Similar structures called cystidia often occur within the hymenium of the Basidiomycota. Many types of cystidia exist, and assessing their presence, shape, and size is often used to verify the identification of a mushroom. [12]

The most important microscopic feature for identification of mushrooms is the spores. Their color, shape, size, attachment, ornamentation, and reaction to chemical tests often can be the crux of an identification. A spore often has a protrusion at one end, called an apiculus, which is the point of attachment to the basidium, termed the apical germ pore, from which the hypha emerges when the spore germinates. [12]

Many species of mushrooms seemingly appear overnight, growing or expanding rapidly. This phenomenon is the source of several common expressions in the English language including "to mushroom" or "mushrooming" (expanding rapidly in size or scope) and "to pop up like a mushroom" (to appear unexpectedly and quickly). In reality, all species of mushrooms take several days to form primordial mushroom fruit bodies, though they do expand rapidly by the absorption of fluids. [ citation needed ]

The cultivated mushroom, as well as the common field mushroom, initially form a minute fruiting body, referred to as the pin stage because of their small size. Slightly expanded, they are called buttons, once again because of the relative size and shape. Once such stages are formed, the mushroom can rapidly pull in water from its mycelium and expand, mainly by inflating preformed cells that took several days to form in the primordia. [ citation needed ]

Similarly, there are other mushrooms, like Parasola plicatilis (formerly Coprinus plicatlis), that grow rapidly overnight and may disappear by late afternoon on a hot day after rainfall. [13] The primordia form at ground level in lawns in humid spaces under the thatch and after heavy rainfall or in dewy conditions balloon to full size in a few hours, release spores, and then collapse. They "mushroom" to full size. [ citation needed ]

Not all mushrooms expand overnight some grow very slowly and add tissue to their fruiting bodies by growing from the edges of the colony or by inserting hyphae. For example, Pleurotus nebrodensis grows slowly, and because of this combined with human collection, it is now critically endangered. [14]

Though mushroom fruiting bodies are short-lived, the underlying mycelium can itself be long-lived and massive. A colony of Armillaria solidipes (formerly known as Armillaria ostoyae) in Malheur National Forest in the United States is estimated to be 2,400 years old, possibly older, and spans an estimated 2,200 acres (8.9 km 2 ). [15] Most of the fungus is underground and in decaying wood or dying tree roots in the form of white mycelia combined with black shoelace-like rhizomorphs that bridge colonized separated woody substrates. [16]


Contents

PIP2 is a part of many cellular signaling pathways, including PIP2 cycle, PI3K signalling, and PI5P metabolism. [4] Recently, it has been found in the nucleus [5] with unknown function.

Cytoskeleton dynamics near membranes Edit

PIP2 regulates the organization, polymerization, and branching of filamentous actin (F-actin) via direct binding to F-actin regulatory proteins. [6]

Endocytosis and exocytosis Edit

The first evidence that indicated phosphoinositides(PIs) (especially PI(4,5)P2) are important during the exocytosis process was in 1990. Emberhard et al. [7] found that the application of PI-specific phospholipase C into digitonin-permeabilized chromaffin cells decreased PI levels, and inhibited calcium-triggered exocytosis. This exocytosis inhibition was preferential for an ATP-dependent stage, indicating PI function was required for secretion. Later studies identified associated proteins necessary during this stage, such as phosphatidylinositol transfer protein , [8] and phosphoinositol-4-monophosphatase 5 kinase type Iγ (PIPKγ) , [9] which mediates PI(4,5)P2 restoration in permeable cell incubation in an ATP-dependent way. In these later studies, PI(4,5)P2 specific antibodies strongly inhibited exocytosis, thus providing direct evidence that PI(4,5)P2 plays a pivotal role during the LDCV (Large dense core vesicle) exocytosis process.

Through the use of PI-specific kinase/phosphatase identification and PI antibody/drug/blocker discovery, the role of PI (especially PI(4,5)P2) in secretion regulation was extensively investigated. Studies utilizing PHPLCδ1 domain over-expression (acting as PI(4,5)P2 buffer or blocker) , [10] PIPKIγ knockout in chromaffin cell [11] and in central nerve system [12] , PIPKIγ knockdown in beta cell lines , [13] and over-expression of membrane-tethered inositol 5-phosphatase domain of synaptojanin 1 , [14] all suggested vesicle (synaptic vesicle and LDCV) secretion were severely impaired after PI(4,5)P2 depletion or blockage. Moreover, some studies [14] [12] [11] showed an impaired/reduced RRP of those vesicles, though the docked vesicle number were not altered [11] after PI(4,5)P2 depletion, indicating a defect at a pre-fusion stage (priming stage). Follow-up studies indicated that PI(4,5)P2 interactions with CAPS, [15] Munc13 [16] and synaptotagmin1 [17] are likely to play a role in this PI(4,5)P2 dependent priming defect.

IP3/DAG pathway [18] Edit

PIP2 functions as an intermediate in the IP3/DAG pathway, which is initiated by ligands binding to G protein-coupled receptors activating the Gq alpha subunit. PtdIns(4,5)P2 is a substrate for hydrolysis by phospholipase C (PLC), a membrane-bound enzyme activated through protein receptors such as α1 adrenergic receptors. PIP2 regulates the function of many membrane proteins and ion channels, such as the M-channel. The products of the PLC catalyzation of PIP2 are inositol 1,4,5-trisphosphate (InsP3 IP3) and diacylglycerol (DAG), both of which function as second messengers. In this cascade, DAG remains on the cell membrane and activates the signal cascade by activating protein kinase C (PKC). PKC in turn activates other cytosolic proteins by phosphorylating them. The effect of PKC could be reversed by phosphatases. IP3 enters the cytoplasm and activates IP3 receptors on the smooth endoplasmic reticulum (ER), which opens calcium channels on the smooth ER, allowing mobilization of calcium ions through specific Ca 2+ channels into the cytosol. Calcium participates in the cascade by activating other proteins.

Docking phospholipids Edit

Class I PI 3-kinases phosphorylate PtdIns(4,5)P2 forming phosphatidylinositol (3,4,5)-trisphosphate (PtdIns(3,4,5)P3) and PtdIns(4,5)P2 can be converted from PtdIns4P. PtdIns4P, PtdIns(3,4,5)P3 and PtdIns(4,5)P2 not only act as substrates for enzymes but also serve as docking phospholipids that bind specific domains that promote the recruitment of proteins to the plasma membrane and subsequent activation of signaling cascades. [19] [20]

  • Examples of proteins activated by PtdIns(3,4,5)P3 are AKT, PDPK1, Btk1.
  • One mechanism for direct effect of PtdIns(4,5)P2 is opening of Na + channels as a minor function in growth hormone release by growth hormone-releasing hormone. [21]

Potassium channels Edit

Inwardly rectifying potassium channels have been shown to require docking of PIP2 for channel activity. [22] [23]

G protein-coupled receptors Edit

PtdIns(4,5)P2 has been shown to stabilize the active states of Class A G protein-coupled receptors (GPCRs) via direct binding, and enhance their selectivity toward certain G proteins. [24]

G protein-coupled receptor kinases Edit

PIP2 has been shown to recruit G protein-coupled receptor kinase 2 (GRK2) to the membrane by binding to the large lobe of GRK2. This stabilizes GRK2 and also orients it in a way that allows for more efficient phosphorylation of the beta adrenergic receptor, a type of GPCR. [25]

Regulation Edit

PIP2 is regulated by many different components. One emerging hypothesis is that PIP2 concentration is maintained locally. Some of the factors involved in PIP2 regulation are: [26]


4.5: Macrofungi - Biology

Fungi are the most diverse organisms on earth and are defined as a eukaryotic, heterotrophic which is devoid of chlorophyll and obtains its nutrients by absorption and reproduces by means of spores [ 1 ]. Large fungi are those that form large fructifications visible without the aid of the microscope and include Basidiomycota and Ascomycota with large observable spore bearing structures [ 2 , 3 ]. Ecologically, macrofungi can be classified into three groups: the saprophytes, the parasites, and the symbiotic (mycorrhizal) species. Most terrestrial fungi are saprobes or mycorrhizal symbionts, but some are pathogens of plants or fungi. Macrofungi fruiting on woody substrate are usually either saprobes or plant pathogens [ 4 , 5 ]. Fungi of various taxonomic groups producing conspicuous sporocarps are collectively known as macrofungi which include “gilled fungi,” “jelly fungi,” 𠇌oral fungi,” “stink fungi,” 𠇋racket fungi,” “puffballs,” “truffles,” and 𠇋irds nest” [ 6 ]. Macrofungal diversity is an important component of the global diversity, particularly community diversity, which is an essential part of fungal diversity [ 7 ]. Mushrooms are widespread in nature and they still remain the earliest form of fungi known to mankind [ 8 ].

Only about 6.7% of the 1.5 million species of fungi estimated in the world have been described and these are mostly in temperate regions. The tropical region which has the highest fungal diversity has not been fully exploited [ 9 ]. Cameroon has a rich biodiversity but it remains poorly unexplored. Termitomyces spp. are widely distributed across the country and form an important source of income for the rural people of Baligham and Ndop plains of the Northwest Region of Cameroon as well as Mbouda in the Western part of the country [ 10 ]. Checklist of macrofungi of Mount Cameroon consisted of 177 species as reported by [ 11 ].

Wild edible mushrooms are one of the most important natural resources on which the people of many nationalities rely and play a key role in nutrition [ 12 ]. Ethnomycology investigates the indigenous knowledge of mushroom utilization and consumption patterns such as in nutrition, medicine, and other uses [ 13 ]. It also investigates the ectomycorrhizal association and ecological benefits of macrofungi (mushrooms) to the forest. In Cameroon, mushrooms are known and consumed in many households, in the country sides and in forest areas [ 13 ]. During the onset of the rainy season when mushrooms are abundant most people in the rural areas collect them from the forest for consumption and sale [ 10 ]. The current rate of bush burning, deforestation, and overexploitation of both timber and nontimber products are threatening mushroom diversity in Cameroon. The use of fungi for food and medicine goes back a long way in human history, but research and documentation of such knowledge are relatively new in Cameroon even though one hundred and seventy-seven species of mushroom were identified in the Mount Cameroon Region [ 11 ]. Based on literature available to us macrofungi diversity in the Awing forest reserve has not been studied and there is no documentation on their ethnomycological knowledge. It is therefore crucial to document the diversity and ethnomycology of macrofungi in the Awing forest reserve and communities. Hence the objective of this study was to investigate the mushroom species richness in the Awing forest reserve with the aim of producing a checklist of macrofungi for the area and also to document the traditional knowledge of mushrooms in the communities surrounding the Awing forest reserve.

2. Materials and Methods 2.1. Study Area

Awing is found in the Northwest Region of Cameroon in West-Central Africa. It is located between latitude 05끑.527′N and longitude 010뀒.122𠌮, with an altitude of 2126 m. Awing has a surface area of about 100 km 2 . The climate is tropical with dry and rainy seasons. It has a humidity of 98% and it is a grass-field area with fertile volcanic soils. The map of the sampled area in the Awing forest reserves for diversity studies is found in Figure 1 while the map of the sampled area in the Awing communities used for ethnomycological studies is shown in Figure 2 .

Awing forest reserves showing the area for diversity studies.

Map of Awing showing communities used for ethnomycological studies.

The latitude, longitude, and elevation for the three plots used for diversity studies are shown in Table 1 .

Location of sample sites used for diversity studies of macrofungi in the Awing forest reserve.

The sampling sites were chosen based on the accessibility of the area and presence of macrofungi [ 6 ]. The field protocol was according to [ 14 ], in which repeated sampling of all macrofungi species present in the sites was done for six months from February to July 2015. Sampling of the macrofungi was carried out using transects of 50 × 20 m in three different plots consisting of high altitude, mid altitude, and low altitude. Photographs of the macrofungi species were taken in situ and macromorphological characters recorded [ 6 , 14 ]. The collection of all macrofungi species was done with care to avoid damage of the sporocarp and they were wrapped in tissue and placed in separate collection bags to avoid spore contamination among the different species of macrofungi. The drying of the macrofungi samples was done using a portable plant drier at 25�ଌ for 2-3 days depending on the texture of the fruiting body. Identification, taxonomic keys, and descriptions were consulted according to [ 15 ]. The samples representing TK1-TK75 have been stored in the Department of Biological Sciences Laboratory, Faculty of Science, The University of Bamenda, Cameroon.

2.3. Ethnomycology Documentation

Those involved in the ethnomycology studies included the aged males and females, traditional practitioners (alternative medicine), and elites of the community. Their consent was gotten before the initiation of discussion and administering of questionnaires. A focus group discussion was carried out and interviews were made accompanied by great participation of the indigenes of the communities. One hundred questionnaires were administered in each of the fifteen villages, followed by a question and answer session where both the informant and researcher asked and answered questions. The questions included informant’s data (which included the name, occupation, migratory history, land tenure, and family size), list of all mushrooms the informant knew, for example, the traditional name, description, time of occurrence, habitat, and its relationship with plants and animals. Informants gave their knowledge of macrofungi and their uses. It was also asked whether the inhabitants attached myths to mushrooms and the informant’s relationship with the forest. A pictorial presentation was also done where the communities identified the mushroom giving their vernacular names and uses. About 250 mushroom pictures obtained from a biodiversity survey from the Awing forest reserve area were presented to the communities members.

3. Results and Discussion 3.1. Species Richness

A total of 75 species of mushrooms in 30 families belonging to 7 Ascomycota and 68 Basidiomycota were identified during the entire period as shown in Table 2 .

Checklist of macrofungi in the Awing forest reserve.

The species richness tends to decrease with increase of altitude with the highest one at the lowermost altitude and the lowest one at the highest altitude. Thirty-six species were collected only from the low altitude, 16 species from the mid altitude, and 16 species from the high altitude. No species was common to all the three altitudes, 1 species was common to both high and low altitudes, and 5 species were common to both the high and mid altitudes, while 1 species was common to both the mid and low altitudes (Figure 3 ).

Species richness across altitude in the Awing forest reserve.

From the focus group discussion and information obtained from the questionnaire and pictorial presentation, it was realized that many people in the Awing communities were familiar with mushroom and its uses as food and medicine and for mythological purposes. The local population commonly calls mushroom “Poh” but specifically “Pohnu” for edible mushroom and “Pohperseh” for poisonous mushrooms. The edible mushroom is usually substituted for animal protein and it is called meat for the poor. No cases of mushroom poisoning were recorded among the people. Some people in the Lake Awing Area did not consume mushroom (mycophobic) but those actively involved in consumption and utilization claimed that it was inherited from their forefathers. The aspect of inheritance is in line with the findings of [ 16 ] that studied the sociocultural and ethnomythological uses of edible and medicinal mushrooms found in the Igala land in Nigeria and [ 13 ] that studied the ethnomycology of edible and medicinal mushroom in the Mount Cameroon Region. Some informants from the communities said that they also consume mushroom because of its nutritive value, as a protein source because they regard it as substitute for meat, some consume mushroom because it is tasteful and for its medicinal value. The communities could distinguish between edible and poisonous mushrooms. The people of the Awing communities claimed that when insects or animals (rabbit, grass cutters, and tortoise) feed on mushrooms they know that they are edible. The people also said that if the mushroom is rubbed on sensitive parts of the body such as the inner part of the elbow and the navel and it itches, then it is poisonous. Moreover, they said brightly coloured mushrooms are mostly poisonous while dull coloured mushrooms are edible. Species commonly used as food and medicine in the Awing communities included Termitomyces titanicus , Laetiporus sulphureus , and Auricularia auricula . This is contrary to the findings of [ 17 ] that noted the consumption of mostly Termitomyces sp., Cantharellus sp., Volvariella sp., Lentinus squarrosulus , and Lactarius spp. in the South of Cameroon. Reference [ 13 ] recorded that species used for ethnomedicine among the Bakweris communities in Mount Cameroon Region belonged to several genera, including Termitomyces , Auricularia , Agaricus , Daldinia, Dictyophora, Pleurotus, Russula, Trametes, Chlorophyllum, and Ganoderma . Species used for ethnomycology among the Awing people belong to several genera including Termitomyces, Laetiporus, and Agaricus while species such as Termitomyces titanicus and Termitomyces microcarpus were found to possess mythological uses (Table 3 ).

Ethnomycological uses of mushrooms in the Awing communities.

This study revealed that mushroom gathering is an important economic activity whose sustenance was threatened by the erosion of the biodiversity. It was found that mushroom harvesting is gender related, being generally regarded as work for women and children this corroborates the findings of [ 18 ] among the Igbo people of Nigeria and [ 13 ] in the Mount Cameroon Region. Pictures of some mushrooms identified by the communities of Awing are shown in Figure 4 .

Some mushrooms in the Awing forest reserve: (a) Auricularia auricular , (b) Laetiporus sulphureus , (c) Ganoderma sp., (d) Auricularia delicata , (e) Cordyceps robertsii , (f) Oudemansiella canarii , (g) Gyrodon merulioides , (h) Ramaria sp., (i) Xylaria ianthinovelutina , (j) Pleurotus ostreatus , (k) Stereum ostrea , (l) Trametes sp., and (m) Geastrum triplex.

(a) (b) (c) (d) (e) (f) (g) (h) (i) (j) (k) (l) (m) 4. Conclusion

The list of macrofungi in this study provides the baseline information needed for the assessment of changes in mushroom biological diversity in the Lake Awing Area. It is an important first step towards producing a checklist of macrofungi in the Lake Awing Area. For the first time in the records of Cameroon, Cordyceps robertsii , the medicinal caterpillar fungus, was identified as a new record for Cameroon. The indigenes of the Awing communities lack ethnomycology knowledge compared to other communities studied in Cameroon. There is increasing interest in the mapping of macrofungi in many areas to obtain the distribution records similar to those already existing for flowering plants. However, unlike plants the identification of macrofungi relies on the collection of fruiting bodies, which in turn is largely dependent on the availability of moisture in most cases. The importance of mushrooms is not only in the ecosystem dynamics but also in human nutrition and health and hence increases the need for the conservation of this nontimber forest product resource. Conservation can be achieved through cultivation, creation, and protection of forest reserve areas and preservation of mushroom habitat. It is therefore necessary to include macrofungi biodiversity conservation in forest management policies in Cameroon.


UTC Scholar

I conducted a survey of the macroscopic fungi within Cloudland Canyon State Park, Dade County, GA that consisted of twenty-three forays from May through December of 2019, and one foray in March 2020. The results of my survey add baseline data to our knowledge of the mushrooms present within the park, allow for the future construction of an All Taxa Biodiversity Index, and allow comparisons to other surveys of fungal diversity in similar areas of the Cumberland Plateau: the Tennessee River Gorge Trust (Starrett 2005), and the Lula Lake Land Trust (De Guzman 2000). My survey resulted in an overall collection of 198 specimens of which 116 were identified. Of the 116 specimens identified, 55 genera and 70 species were recorded. Specimens collected for this survey will be accessioned in the UTC Museum of Natural History - Fungi, and images and metadata will be uploaded to MycoPortal. My research objective was to contribute to the knowledge of the macrofungi of the southern Cumberland Uplands. The aim of the present study was to add species to the lists of those macrofungi known to occur within the bounds of the large, nearly contiguous public and private conservation lands of The Tennessee River Gorge, the Lula Lake Land Trust, and Cloudland Canyon State Park. These three areas are similar geologically, geographically, floristically, and have a rich, shared cultural history. The Jaccard's Index of Similarity was utilized in comparing the similarities of macrofungi within Cloudland Canyon State Park, the Tennessee River Gorge Trust, and Lula Lake Land Trust.

Acknowledgments

I would like to extend my gratitude to Brad Gibson for allowing me to forage in this place that has become so significant to me and for managing it well. This place has brought me much peace and discovery amidst a busy season of my life, and for that I am thankful. I would also like to extend my thanks to my thesis director and friend, Dr. J. Hill Craddock. Without him taking a chance on me, this vast and beautiful world of fungi would have never meant as much to me as it does now. His whim and knowledge made the hard parts bearable and magnified the fun ones. Thank you to my committee members, Dr. Jennifer Boyd and Dr. David Aborn, for joining me in this moment and inviting me to view it through a different lens. I owe an immense amount of thanks to my parents and friends, all of whom stuck with me talking about mushrooms and were truly excited for me. Lastly, thank you to those who joined me on forages and were willing to learn and help me: Dalton Strike, Ashley Carpenter, Molly Stelling, Trent Sims, Nicole Elmore, Jacob Goldsmith, Leanah Chestnut, Bella Horrocks, Savannah Sarwar, Hill Craddock, and Paola Zannini. You all made the load of writing a thesis lighter in these shared moments of discovery.

Degree

B. A. An honors thesis submitted to the faculty of the University of Tennessee at Chattanooga in partial fulfillment of the requirements of the degree of Bachelor of Arts.