Lecture 04: Naming and Classification of Fungi

Introduction

Although it is something that we give little thought to, the naming and classification of objects play an important role in society. Imagine the problems that we would have if we were not given names and if we did not name our cities, streets or pets. Now imagine if we did not classify the books that are in our libraries, tools in hardware stores, video tapes in video stores, etc. Without names for objects and a system to classify them, it would be impossible or at least very difficult to organize, categorize and find information for which we are searching. It should not be a surprise then that there is a discipline in biology called taxonomy that studies the classification and naming of organisms. Presently, there are approximately 1.5 million species of organisms that have been described by biologists, and each year thousands of new species are described by science. It is estimated that the total number of species may range from 5 to 30 million. With such a rich diversity of life on this planet, naming and classifying these many organisms would be a difficult task, to say the least. However, such a system must be in place if our knowledge in biology is to advance.

The job of the taxonomist is a difficult one. The lay public thinks of them as being snobbish because they refer to   organisms with "fancy scientific names". The reputation of the taxonomist is often not regarded in high esteem even among scientist because of its descriptive rather than experimental nature. Christensen (1961), a well-respected mycologist-plant pathologist, describes taxonomist as "those who spend their time putting living things into pigeonholes, a low but necessary form of scientific endeavor". The lay person often wonders why taxonomist must refer to onions as Alium cepa or leopards as Panthera pardus. However, they are not using these names just to impress you or to make themselves feel more superior. Instead, they are merely identifying organisms by their scientific or species name. The species name is called a binomial, which consists of two parts. Binomial literally means two names. The first part is the genus (plural: genera) and the second is the specific epithet. For example, the species name for the human animal is Homo sapien. When written, the species name is emphasized by italicizing or underlining, i.e. Homo sapien the binomial. The genus is always capitalized and the specific epithet is always in lower case, even when it is a proper name, for example Agra schwarzeneggeri is named after Arnold Schwarzenegger, but the specific epithet is not capitalized. Each known species, once it has been described by science, is given a unique species name.  Another words, there is only one valid scientific name that is unique to a species regardless of where it may be found. The reason for this is there is an international agreement among taxonomist for different groups of organisms that a species name must refer to only one species. This differs from how society would name an organism. For example, when we name our pets or children, the selected names are not unique. After all, how likely is it that anybody at this time would come up with an original, unique name. In taxonomy those types of names would be referred to as common names, which have been given to many species of organisms. It is not unusual for several common names to be applied to a single species and often a common name may be applied to several species. Although many well-known species of organisms have common names, they are not officially used in science because of the confusion that they may cause.  For example, the common name "Spanish Moss" is applied to at least two species of unrelated organisms in North America, Ramalina menziesii (Fig. 1) and Tillandsia usenoides (Fig. 2). The former is a lichen, which represents a symbiotic relationship between a fungus and an alga and the latter is a flowering plant. 

Not only is this an example of a common name that applies to two different, unrelated species, but also one, which is misleading since neither species is a moss, nor is it Spanish. Conversely, Plantago major, a widespread weed, in Europe, is sometimes called "broad leafed plantain", but it also has at least 45 other English common names. Because the European countries speak a number of different languages, there are also 11 French names, 75 Dutch names, 106 German names and many more in other languages (Stern, et al, 1997). Many of these common names may also apply to other unrelated plants. Nevertheless, because common names were the first names that many organisms received and are the names that the general population uses, it is unlikely that they will going away anytime soon and be replaced by binomial names in daily usage.

Development of the Binomial System and Classification Schemes

A History of Taxonomy

Although the usage of binomials seems to be a simple enough concept, its development took well over a thousand years. In summarizing the events that led to the binomial system, we will concern ourselves mainly with events having to do with plants because fungi as well as algae and bacteria were classified as plants until Whitaker (1969).

The first names applied to plants were undoubtedly "local" common names. Because such names may be restricted to small communities, another nearby community would likely have a different common name for the same plant. This seems to be the likely mechanism by which numerous common names may have been applied to a single plant.

One of the earliest recorded attempts, in the Western World, at classifying plants was in the 3rd Century B.C., by Theophrastus who classified approximately 500 species of plants into three categories: trees, shrubs and herbs, and utilized leaf characteristics and other vaguer features in distinguishing between different species. If this method of classifying plants had continued in this matter, the progress of plant classification would undoubtedly be further along today. However, a slight detour occurred for over a thousand years.

A more applied scheme of classifying plants was devised around the 2nd Century A.D. Dioscorides, a Greek physician, classified plants according to their medicinal usage. He illustrated and described 600 species of plants in his book, De Materia Medica. His books, which were highly respected, were frequently reproduced by hand copying and became known as herbals. Unfortunately, there is such a thing as being too highly esteemed. The herbals became the sole source of medicinal information as well as plant classification between 400 and 1400 A.D. (Why is classifying a plant according to its medicinal use not as desirable as using characteristics of plant structure, as Theophrastus had done?) It was considered heresy to speak against or to question information in the herbals, which resulted in few new ideas during this period of time. This was the period referred to as the Dark and Middle Ages, in Europe, and for a thousand years the advancement of knowledge in science slowed.

It would not be until the 15th Century that a proliferation of knowledge would come about. This was, in part, due to new ideas that came about, but also had to do with the invention of the printing press by Johannes Gutenberg, which made the mass reproduction of books possible. Prior to this invention, books were hand written. Thus, few books were available. This was also when the herbalists began to classify plant species according to their usage. This differed from the herbals in that the herbalist now classified plants not only for their medicinal usage, but also by their usage as food, clothing, as well as other applications. Included among these plants were also usages of some species of fungi.

By this time botanists were once again classifying plants in the matter of Theophrastus, i.e. comparing the characteristics of different plant structures. Andrea Caesalpino (1519–1603) was on of the earliest, using floral parts as well as fruit, seed characteristics, as well as other structures that Theophrastus did not use. During this period, it was also realized by Jean and Gaspard Bauhin (1541–1631; 1560–1624) that many plants had many different names and introduced the concept of synonymy and in their publications listed names of plants that referred to the same species. Although botanist continue to give credit to Linnaeus as the originator of the binomial system of naming species, it was actually Gaspar Bauhin that first used binomial names in his Pinax Theatri Botanici, nearly two centuries before Linnaeus. However, usage of binomials did not catch on at that time. Instead what became more popular was the use of phrase name in naming plants. A phrase name both described as well as named plant species and even credited these names to the authority, the person who named the plant, for each species. Also, the use of a phrase name usually consisted of no more than 12 words. However, as in the case of the common names, there was still more than one phrase name for each species. Different phrase names applied to the same species often differed by only a few adjectives.

 In 1700 Joseph Pitton de Tournefort (1656–1708) originated the concept of phrase names that began with the genus of the species followed by a short Latin description.  For example, the phrase name for the spearmint plant was Mentha floribus spicatis, foliis oblongis serratis. Mentha is the genus name followed by a brief Latin description. By this time, Latin had become widespread in schools and universities throughout Europe. Thus, phrase names were written in Latin for ease of communication. In English, the Latin phrase name of the spearmint plant would translate to "mint with flowers in a spike, leaves oblong and toothed". For obvious reasons, a phrase name was not practical for naming organisms. Can you imagine having to repeat the above scientific name each time reference was made to spearmint? However, their use persisted until the reintroduction of binomials, in 1753, by Linnaeus..

Carolus Linnaeus was credited with originating the binomial species names in his 1753 publication of Species Plantarum and he also "popularized" botany at this time. Less has been said of how he popularized it. At the time of Linnaeus, it was realized that plants, like animals, reproduced sexually. Using this knowledge, Linnaeus published in 1735, his Systema Naturae, on his sexual classification of plants. His classification scheme included counting the number of male (stamens) and female (pistils) reproductive organs that were present in flowers, which seems innocent enough.  However, it was the matter in which Linnaeus represented sexual reproduction in plants that became somewhat controversial. On the sexuality of plants, Linnaeus wrote the following:

His metaphors concerning reproduction in plants, using human analogies proved to be too explicit for the moral values of his time. An excellent example of how Linnaeus used the characteristics of the flower to distinguish between the different classes can be found at http://courses.eeb.utoronto.ca/eeb337/B_How/307b1Linn.html:

Characters of the Classes

• Public Marriages (Flowers visible to everyone)

  • In one Bed (Husband and wife have the same bed)

    • Without Affinity (Husbands not related to each other)

      • With Equality (All the males of equal rank)

        • Varying numbers of males:

 While this sexual system of classification proved to be easy to learn and made it possible for the non-botanist to identify plants, it also invited much criticism, some from his contemporaries. They included such comments as:

 Siegesbeck was his biggest critics and made many other such comments. However, Linnaeus may have gotten the last laugh when he named an ugly, smelly, little weed Siegesbeckia orientalis, after Siegesbeck. While controversial, Linnaeus' controversy of his sexual system is largely forgotten and he is best remembered for his reintroduction of the the binomial system first originated by Gasbar Bauhin.

Linnaeus' Species Plantarum, two-volume publication was an ambitious effort to name and classify all of the known plants during Linnaeus' time. As with most taxonomist of his time, he actually used phrase names in recording the different species of plants that he recognized. However, Linnaeus added a shorthand notation for each species. In the left margin of each phrase name, he wrote a word, which, when combined with the genus, formed a convenient abbreviation for the phrase name. For example, "spicatis" was added to the margin of the phrase name for spearmint and "piperata" was added to peppermint (Fig. 4). The abbreviation for these species could then be referred to as Mentha spicatis and Mentha piperata, respectively. Other workers soon followed Linnaeus in recording species names in this fashion and eventually phrase names were replaced by binomials.

Although Linnaeus has been credited with inventing the usage of binomials for species names, he did credit Gaspar Bauhin with originating this system.

Today, all species names are binomials, and if cited formally, the name of the author who described the species or an abbreviation of the author's name is placed after the binomial, e.g. Mentha spicatis L. The "L." in this case credits Linnaeus as the person who first described this species. There are also variations as to how a person is credited after a species name. For example, in the supermarket mushroom, Agaricus bisporus (Lange) Imbach, Lange, whose name is in parentheses, was the original person who described the species. The parentheses indicate that the species was originally described in another genus. In this case the genus was Psalliota. Imbach, whose name appears after Lange, is credited with making the change from Psalliota to Agaricus. Therefore, if we wanted to find all of the literature that is available on this species, we would have to look under both Psalliota bisporus Lange and Agaricus bisporus (Lange) Imbach. The idea of citing an author's name to credit that person with the discovery of the species and/or the person changing the name of the species also did not originate with Linnaeus. Botanist had long attributed authors' names to species so that a history of changes that occurred in the species names could be followed. Also, it was a matter of giving credit, as well as the blame, for the changes that were made to a species name. There are various reasons for changing the name of a species and such notations allow us to trace these changes. Commonly, a name has to be changed because two species have the same name. Because of this and numerous other problems that can arise, there must be rules that govern the procedures for the naming of species as well as provisions for problems that may arise.

In discussing binomials, some mention has already been made concerning certain practices in the naming of species and their citations, i.e. the name is in Latin, it is underlined or italicized with the author of the species following the epithet. Although it is not obvious, a little thought on the subject should tell us that unless there is agreement, among taxonomists, concerning the method by which plants are named and classified, we can still expect confusion to reign. However, it would be over a hundred years before such an agreement was made. In 1867, 150 botanists, mostly from Europe and America, met in the First International Botanical Congress, to standardize the rules and regulations that would be used when plants are named. The rules adopted by these botanists are known as the International Code of Nomenclature for Fungi, Algae and Plants ( ICN), formerly the International Code of Botanical Nomenclature, and has been adopted by botanists throughout the world. The ICN not only describes how species are named, but also when names must be changed, what names are unacceptable, how to validly publish a new species that you have just discovered and many other topics having to do with taxonomy of plants. The ICN also governs the taxonomy of algae, cyanobacteria [other bacteria are governed by the International Code of Nomenclature of Bacteria] and fungi. Zoologist later adopted The International Code of Zoological Nomenclature that governs the naming of animals.

One of the important rules that was agreed upon after many years of debates was that the "starting date" of plant names should begin with 1753. The reason for this selection was that this was the year that Species Plantarum, which included the binomial names of just about all of the known plants at that time, was published. Species names published earlier than 1753 and not recorded in Species Plantarum were not recognized as valid names. However, Linnaeus included very few species names of fungi and a different starting point for species names of fungi was originally used. There were originally two starting points for fungi: Christiaan H. Persoon's Synopsis Methodica Fungorum issued in 1801 and Elias Magnus Fries' Systema Fungorum, first volume (of three) issued in 1821. Persoon was the starting point for rust and smut fungi and Fries was the starting point for the remaining fungi, especially fleshy fungi, i.e. those with fruiting bodies. In the 1987 International Botanical Congress, this all changed. It was decided that the starting date for valid species names of fungi would be moved to 1753. However, species recognized in Synopsis Methodica Fungorum and Systema Fungorum were sanctioned, which means that even if there was an earlier name for a species published in these tomes, they would still have priority in spite of the fact that there was an earlier species name. Confused?

This hierarch of categories , above, goes from one that is the most inclusive, i.e. Kingdom, to one that is most exclusive, i.e. species.

Even with the progress that have occurred in molecular biology, there is still disagreement even at the broadest categories of our classification schemes. Intuitively, kingdoms would appear to straightforward, but this is not the case. As more is learned from molecular studies of various organisms, this has only caused more disagreement as to the classification scheme to use at the kingdom level.

The Concepts of Kingdoms

The kingdom in an introductory biology course is usually our broadest category and in the earliest classification, only two were recognized, Plants and Animals. Plants were then characterized as organisms that lack motility, and do not consume food while animals have the ability to move and eat their food. Even today, the lay public still categorize life in the two kingdoms, and for most organisms that we are in daily contact with this is a workable system. We know that pines, ferns and mosses are different types of plants and that dogs, birds and fishes are different categories of animals. However, most microscopic organisms do not fit comfortably into either kingdoms. Fungi, for example, were placed into the Plant Kingdom because they lacked motility and their cells were surrounded by a rigid cell wall, but unlike plants they cannot photosynthesize. The genus Euglena (Fig. 3) is one of many examples that have been classified as both a plant and an animal. It is a unicellular organism found in fresh water and can swim through the water with a "hair-like tail" called a flagellum (plural: flagella). A groove or a gullet is present in the cell that allows it to ingest food. However, Euglena also has chloroplasts and, like plants, can make its own food through photosynthesis. Because Euglena possess both plant and animal characteristics, it was classified as both a plant and an animal.  Despite these problems, this two kingdom system was used until Whitaker (1969) proposed that organisms be classified into five kingdoms: Monera (=Bacteria), Protista (=Mostly Algae and Protozoans), Plantae (=Plants), Mycetae (=Fungi) and Animalia (=Animals). Until recently, Whitaker's five kingdoms was the system of classification of organisms used in textbooks since the early 1970's. 

Although Whitaker was credited with this system of classification, most of the additional kingdoms that he recognized did not originate with him. It had long been accepted, in biology, that many organisms did not fit comfortably into a two kingdom system. A third kingdom, Protoctista, was first proposed by the German biologists J. Hogg and Ernst Haeckel, in 1860. This kingdom included organisms such as fungi, bacteria, algae and protozoans that were characterized by having simple cellular organization and not producing complex tissues, and also often had a combination of plant and animal characteristics. However, it was obvious that these organisms were not closely related and did not form a natural grouping. A modification of this system was proposed by Herbert Copeland, in 1938. He removed bacteria from Protoctista and erected a fourth kingdom that he called Monera. From here, the modification from Copeland's classification was not a big leap to Whitaker's, which differed only in the removal of the fungi from Protoctista and placing them in their own kingdom (=Mycetae). It should also be noted that while a five kingdom system of classifying organisms has been used in most text books, for more than 25 years, the concepts of Protoctista, now Protista in most textbooks, and Mycetae have changed during this period of time. With these changes in concepts, other classification schemes have been proposed. Some schemes have expanded the number of kingdoms to six, and eight and one has even reduced the number to three. Thus, there is still not agreement among scientist as to the number of kingdoms that should be recognized in the classification of organisms, and it is probable that there never will be.

The lack of agreement on a single classification scheme does not mean that one system is necessarily better than another. Only that they are different. One point to keep in mind concerning the various classification schemes is that they are man-made and are only our concepts of how organisms should be classified. The organisms, themselves, have not been changed one iota as a result of these changes in classification, even though this is the impression that we are occasionally left with. One example that might amuse you is the change in policy of the Hawai‘i State Plant Quarantine Department, with respect to a group of organisms commonly referred to as "blue-green algae". Plant quarantine's import policy of this group of "algae" was to treat them as algae. However, after they had learned that blue-green algae were reclassified, and placed in the Kingdom Monera, which met that they were now interpreted as being closely related to bacteria rather than algae. All of a sudden, the import policy regarding blue-green algae also changed and are now treated as bacteria. Blue-green algae have probably undergone relatively few changes in over a billion years, but yet the change in policy by the State of Hawai‘i would seem to indicate that there was a drastic change in these organisms simply because they have been reclassified into a different kingdom. 

Review of Kingdom and How to Construct a Dichotomous Key

There are commonly five phyla that are recognized in the traditional Kingdoms of Fungi (More are currently recognized, but we will use this more traditional classification scheme):

1. Chytridiomycota: Flagellated zoospores and gametes

2. Zygomycota: Zygospore supported by suspensors on either side.

3. Ascomycota: Ascospores borne with asci.

4. Basidiomycota: Basidiospores borne on sterigmata of basidia

5. Deuteromycota: Sexual stage absent, reproduction by conidia borne on conidiophores.

Dichotomous keys are a means of identifying organisms without the aid of a teacher. This was one of the contributions for which Linnaeus was recognized. When constructing such a key, two choices are offered and the student selects one of the choices based on which choice seems to best fit the organism being identified. Let construct such a key for the five phyla of Fungi above:

A. Flagellated stages present, mostly aquatic, thallus unicellular to mycelial. If

mycelial, coenocytic….….........................................Chytridiomycota

AA. Flagellated stages absent, usually not aquatic, Thallus mycelial or yeast or

both....................................................................................................B

B. Mycelium usually coenocytic, Sexual spore, zygospore, asexual spores

borne in sporangia…..................................……Zygomycota

BB. Thallus mycelial, if present, septate, or yeast, or dimorphic. Sexual spore if

present, not zygospore, asexual spores conidia….................. C

C. Sexual spores ascospores, borne in asci, either on or in fruiting body

or naked..................................................................... Ascomycota

CC. As above, but sexual spores not ascospores borne in asci....D

D. Sexual spores basidiospores, born on

basidia......................................................Basidiomycota

DD. Sexual stage mostly absent, reproduction asexual by

conidia.....................................................Deuteromycota

Usually, after you arrive at a name, the description of the organisms is described after the key. Just as there are different groups of plants, ex., ferns, mosses, conifers, flowering plants, etc., there are different groups of fungi. A brief description will be given for each of the phylum below.

Phylum: Chytridiomycota (Figs. 4-6): Recently, this phylum has been reclassified into several phyla, but we will use the older, more traditional approach to simplify classification in this introduction to the fungi. Using this classification, this will be the only phylum of aquatic fungi. Fungi that have swimming zoospores and gametes. Zoospores are swimming spores that function in asexual reproduction and gametes are sexual spores that reproduce to give us different genetic recombination. Examples of gametes that occur in animals would be sperms and eggs. These stages in the Chytridiomycota are propelled by a single hair-like appendage, attached to the posterior of the cell that is referred to as a flagellum. Members may be unicellular, composed of a single zoosporangium or have short hyphal to extensive mycelial growth that is coenocytic.

Figures 3-5: Zoospore with single posterior flagellum (left), male and female gametangia (middle) and release of male and female gametes (right). Note female gametes are several times larger than males.

Terrestrial Fungi

The remaining phyla of fungi are mostly terrestrial and without a flagellated stage and are classified mostly according to their mode of sexual reproduction.

Phylum: Zygomycota: This phylum is defined by its thick walled zygosporangium that encloses a zygospore (Fig. 7) within, during sexual reproduction. Oftentimes, it is just referred to as the latter. This was before it was realized that it was actually borne within a zygosporangium and the usage of this term is still presently utilized. Mycelium is always present and is always coenocytic.

Phylum Ascomycota: This phylum is defined by production of asci (sing.=ascus) and ascospores (Fig. 8a) produced during sexual reproduction that may be formed on a complex fruiting body referred to as an ascocarp (Fig. 8b-c) that is composed of septate mycelium. Ascocarps may be variable in shape, size and color. Members may also be unicellular. Unicellular members are referred to yeast and reproduce asexually by the process of budding or fission (Fig 9a-b). A budding yeast has a predetermined point where a new cell is blown out, much like a balloon, until it is approximately the size of the original. The nucleus divides and one of the two nuclei migrates into the new cell and the bridge between the two cells is severed, and the two cells separate. The process of fission by comparison is simpler. When the yeast cells attains its optimum size, the nucleus divides and migrate to opposite ends of the cell and a cell wall is laid down, dividing the cell in two. 

Phylum Basidiomycota: This phylum is similar to the Ascomycota in that it also has septate mycelial members and yeast as well. It differs in that during sexual reproduction, basidia and basidiospores (Fig. 10a-b) are produced instead of asci and ascospores. These structures may be borne on fruiting bodies such as the gills of mushrooms (Fig. 10c).

Phylum Deuteromycota: This phylum is defined by sexual reproduction being lost during the evolution of most of its members. However, this is not strictly true. Some species have the ability to reproduce sexually, but do so only in a narrow range of environmental conditions. For that reason sexual reproduction in some species are rarely seen, but when they are discovered, they are usually observed to produce asci and ascospores and thus are reclassified as Ascomycota. Less often they are observed to produce basidia and basidiospores and reclassified as Basidiomycota. Reproduction in this phylum is mostly by asexual spores called conidia that are borne on modified hyphae called conidiophores. Some examples can be seen in Figs. 11-13.

Figures 11-13: 11. Trichocladium conidia borne directly on hyphae. 12. Periconia cluster of conidia borne at tip of conidiophore. 13. Penicillium conidia borne on branched conidiophores.

There are also two additional phyla I would like to mention that were once thought to be "Fungi", but are longer classified as such. The phyla Oomycota (Figs. 14-16) and the Myxomycota (Fig. xx-xx). 

Phylum Oomycota:  The phylum is very fungal in appearance and have most of the fungal characteristics, but does not have a chitinous cell wall, a significant feature that excludes it from being classified as a fungus. They are commonly called "water molds, and are characterized by the large oogonia that contain eggs (Fig. 14). Note that the antheridium (Fig. 14), the male tube containing the sperm nuclei is attached to the oogonium, making possible direct injection of sperm nuclei to the eggs within the oogonium. They have short hyphae to extensive mycelium that is coenocytic. Asexual reproduction also commonly occurs by zoospores (Fig. 15) borne in zoosporangium (Fig. 16). The Oomycota causes some of the more serious plant diseases, some that are of historical significance we will cover. 

Phylum Myxomycota: This phylum is commonly called the the slime molds because of its plasmodium stage, a large single celled structure that contains thousands of nuclei. Although, they were once classified as Fungi, slime molds are not at all fungal like in its appearance. They lack mycelium, do not have cell walls and do not absorb their food. If you viewed the "Rotten World About Us" video, you may recall that slime molds are called the "mold that walks" because they have an amoeba (Fig. 17) and plasmodium (Fig. 18) stage stages that are mobile as they seek food that is consumed through the process of phagocytosis. Probably the only reason that they were classified as fungi is because they reproduce with spores in sporangia (Figs. 19).

Are Classification Schemes and The Rules That Govern the Naming of Plants, Algae and Fungi Really Necessary? The Curtis Gates Lloyd Story.

In reading and hearing about naming and classification and naming of plants, algae and fungi, you can see then that there is not always agreement in the classification of organisms and revisions are constantly being made, based on the regulations in the naming of species as well as other categories in the hierarchy of classification. Furthermore, there are few botanists that know, in detail, or even understand what is in the ICBN. You may even wonder if all this is really necessary. Most botanists, in fact, do agree that we cannot do without these rules and regulations, but there are also many who wonder why species names must change just because it conflicts with the ICBN and question the need for such a document, or even the existence of taxonomy as a discipline within biology. One of the most well-known critics of the ICBN was Curtis Gates Lloyd (Fig. 20). The story of his opinion on this subject and what he did about it makes for another amusing story.

Lloyd was one of the most famous, or infamous mycologists depending on how you feel about this person. Curtis Lloyd was actually not a mycologist by profession. In his first career, he and his two brothers ran a successful wholesale pharmaceutical company called Lloyd Brothers. The company was so successful that by 1917 Lloyd retired while still relatively young and hired a replacement to take charge of his department of the pharmaceutical company so that he could pursue his mycological interest. His interest in mycology came about through his interactions with Andrew Price Morgan, a well known mycologist, of the late 19th and early part of the 20th. Century. It was from Morgan that he learned about and developed his interest in the fungal group commonly referred to as Gasteromycetes, e.g. puffballs.

Although Lloyd eventually became an excellent taxonomist of fungi, he disagreed strongly about the placement of authors' names after the species that they had described or had changed. He felt that this was the main cause for proliferation of species names as well as the large number of name changes and hasty publications. Lloyd believed that this was a means by which "name jugglers" sought to immortalize their own names. Because of his belief, he refused to follow journals' requirement of placing the authors, of species, after binomials, which resulted in the rejection of his manuscripts for publication. However, his manuscripts were published in Mycological Notes, a journal he started, and published between 1898-1925. Although, this journal only published seven volumes and was discontinued after his death, it was a well respected journal with many important publications, and can be found in most university libraries. In addition to his studies on fungi, it was not uncommon for Lloyd to lampoon the practice of "name juggling" in Mycological Notes. Nevertheless, after a number of years, he found that he had not only described a large number of new species, but also had to "juggle" a number of species names as well. One of his notorious acts was his creation of the fictitious Professor N.J. McGinty of Pumkinville Polymorphic Institute, whose name was cited as the authors of species that he described, which included the fictitious Lycoperdon anthropomorphous, as well as for name changes that he had made. Although he would never admit it, the creation of McGinty may have been his way of saying that he understood the necessity for name juggling. However, his criticism of name juggling would continue for the rest of his life. In what was possibly his last "poke" at the mycological establishment, four years before his death, Lloyd prepared his own tombstone, with the following epitaph:

This was a monument erected to himself in the cemetery at Crittenden, Kentucky, which he said "was intended as a burlesque on tombstones in general and a satire on mycologists who have passed".

Many of Lloyd's collections were obtained through requests by mail and in returned he offered his journal free to the senders of collections. His contributions to mycology were many. During his lifetime, he accumulated a rather large herbarium of fungi which is still maintained today as is his library, which has many rare mycological books. His herbarium was composed only of large fungi because he did not like to use a microscope and believed in distinguishing species with only what was visible to the naked eye. He was the stereotypical scientist. He was never married, except to his work. He didn't have a house. He maintained bachelor's quarters in his museum.

Terms and Concepts for Classification and Naming of Fungi

Ascospores: Sexual spores produced in an ascus (pl. = asci) characteristic of phylum Ascomycota.

Basidiospores: Sexual spores produced on a basium (pl.= basidia) characteristic of phylum Basidiomycota.

Binomial: A species name composed of two parts, the genus and specific epithet.

Common name: Name given to a species by local community. Name given to species not governed by rules and may composed of any number of words and may be of any language.

Deuteromycota: Phylum that does not have sexual stage and reproduce asexual spores only.

Doctrine of Signatures: The belief system that the appearance of a plant serves as a "signature" that determines its use by people.

Phrase name: A Latin name composed of the genus followed by a description of 12 words or less.

Herbals: Books with descriptions of plants and their medicinal use.

Herbalists: People who have expertise in the utilization of plants as a means of medicine, textile, food, etc.

Hierarchy of Classification: The utilization of characteristics from the most inclusive to the most exclusive. This system first utilized by Linnaeus enabled non-botanist to identify plants without the aide of a teacher.

Flagellum (pl. = flagella): Hair-like structure that enable Chytridiomycota spores to swim.

International Rules of Nomenclature for Fungi, Algae and Plants: Rules governing the naming of plants, algae and fungi.

Kingdom: The broadest, most inclusive taxonomic classification level into which organisms are grouped. We will only be concerned with the Kingdom Fungi at this level.

Linnaeus: Botanist who popularized botany, developed a system of classification in plants that permitted the identification of plants without a teacher, credited for the use of binomials, hierarchy classification of organisms, and for recording all of the known species of known plants in his Species Plantarum, published in 1753.

Phylum: The taxonomic level following kingdom. We will only use this level to define the different kinds of fungi.

Taxonomy: That part of biology that studies the naming and classification of organisms.

Zygospore or Zygosporangium: Sexual spore characteristic of phylum Zygomycota. Zygospore is actually produced within zygosporangium. However, you will be allowed to use either terms to define the phylum.

Learning Goals Questions

1. What is taxonomy?

2. Why is taxonomy and classification important in everyday life?

3. Why do we use scientific names in biology rather than common names?

4. What is a binomial?

5. Why was it important that hunter gatherer societies named and classified plants?

6. What are herbals and herbalists? How do they differ?

7. What is the Christian definition of the Doctrine of Signatures?

8. What is a phrase name and what was the problem with using them?

9. Why was Latin once used to name and describe species and why do we no longer use it?

10. Who is Linnaeus and what are his contributions to taxonomy/botany?

11. Linnaeus used a “hierarchy” in classification. What is that?

12. What is the International Code of Nomenclature for Fungi, Algae and Plants and what is its function?

13. What are the different phyla of Fungi and what are the main characteristics of each phylum?

14. Who is Curtis Gates Lloyd?

Literature Cited:

Alexopoulos, C. J., C. W. Mims, and M. Blackwell. 1996. Introductory Mycology. John Wiley & Sons, Inc.

Hibbett, D. S. 2006. A Phylogenetic overview of the Agaricomycotina. Mycologia 98: 917-925

Stern, K. R., S. Jansky, and J. E. Bidlack. 1997. Introductory Plant Biology. McGraw Hill.

Webster, UJ. and R. W. S. Weber. 2007. Introduction to Fungi. Cambridge University Press, New York.

Whittaker, R. H. 1969. New concepts of kingdoms or organisms. Evolutionary relations are better represented by new classifications than by the traditional two kingdoms. Science 163: 150–60.