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BIO 341 Lecture Topic 2: Outline of fungal classifications, "Lower fungi," "Higher fungi" (emphasis on basic diagnostic characteristics of divisions, subdivisions and classes; discussions of the nature of spores, sporangia, asci and basidia).

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Corresponding Readings:
  • Moore-Landecker preface, chapter 2
 Lecture Summary:
Dr. Szaniszlo tied this topic into the last one by reviewing some of the concepts of the first - namely his introduction of the definition of a fungus.  He extended the definition with a discussion about the most commonly described vegetative manifestation of the fungi, the mycelium.  The growth forms of the fungi as well as their modes of nutrition are compared and briefly described.  

With the seemingly striking similarities in the physical nature of fungi and plants (sedentary, propagation and growth models), the lecture turns towards making the distinction between them.  This usually provides an idea place to break in the topic, and the lecture concluded with a history of classification systems, discussion of the 5 kingdoms, and the place of fungi within the current scheme of living organisms. 

 End Lecture One (1/20/2000)

The significance of the classification system was discussed as well as its importance to the student in mastering certain concepts and doing well in the class.  The discussion of early classification systems segued into the topic of the Phycomycetes, one of the classes (no longer used today) included in an early classification scheme.  Dr. Szaniszlo elucidated some primary characteristics that led to the classification of certain organisms into the class - relating too the transition of this classification based on morphology to a classification based increasingly on biochemical and molecular data which has led to the current system that he provided as a handout.  

(Note: the handout will be very useful throughout the semester in learning the classification schemes and understanding how the organisms you study relate to each other.  If the material seems daunting, don't panic - you don't have to learn everything verbatim - use a highlighter to mark only the phylum, classes, orders, etc. that Dr. Szaniszlo emphasizes in class.)

End Lecture Two (1/25/2000)

(Summary Forthcoming)

End Lecture Three (1/27/200)

At the beginning of the lecture, Dr. Szaniszlo reiterated the basic classification system.    In discussing the Ascomycota, he reemphasized the differences between the Hemiascomycotina and the Euascomycotina, paying attention to the variety of multihyphal aggregates formed by the latter.  Note these structures and their associated classes - you will see that they have appeared in past tests.  

Of particular interest, he showed a phylogenetic tree showing the relative positions of the various phyla based on 18S rRNA sequencing.  It confirmed the classification scheme developed over years by researchers based primarily on morphological observation alone.

The professor began his discussion of the Basidiomycota with the differences in the subphyla Heterobasidiomycotina and the Holobasidiomycotina.  Primarily the main differentiating characteristic is the production of basidia in Heterobasidiomycotina from a unique spore called the teliospore.  This  spore is dikaryotic -- and prior to germination becomes diploid by karyogamy of two mating nuclei within the spore.  Dr. Szaniszlo concluded the remainder of the lecture by showing pictures of various Holobasidiomycotina.

End Lecture Four (2/1/2000)

As in his previous lectures, Dr. Szaniszlo begins with reviewing the material he has covered previously.  Remember that that up until now, he has covered three major phyla of the Kingdom Fungi, Chytridiomycota, Ascomycota, and Basidiomycota.  Molecular data from gene sequencing confirms that there is a definite basis for the cytological and morphological division of the higher Fungi in Ascomycota and Basidiomycota.  This leads into the professor's discussion of the form phyla, Fungi Imperfecti (Deuteromycota). 

Some time is spent explaining the reasoning for this classification.   There needed to be a division for those fungal organisms exhibiting no sexual cycle.  This "artificial" grouping exhibited the same hierarchy of names as other organisms, save that the prefix, "form" was appended.  e.g. - "form-species, form-genera, form-families, form-orders, etc."  This division arose from a more practical need and provides a means of providing identification, classification, and basic nomenclature of the the organism.  Historically, it has provided a name to reference these anamorphic (asexual) species.  As teleomorphic (sexual) forms of the species were discovered, new names were given to the species -- and as Dr. Szaniszlo showed during lecture, there are many examples of species with two or more names, one or several for each form of the species.

With sequencing, the majority of Fungi Imperfecti are found to have Ascomycetous character.  In fact, when the sexual form of the fungi are discovered, the fungi are found to be Ascomycota.

Some time was spent explaining why many fungi were/are discovered in their asexual forms.  Dr. Szaniszlo takes that opportunity to explain the concept of homothallism (self-fertile) and heterothallism (self-sterile).  Remember that most fungi are heterothallic and thus are self-sterile, requiring the opposing mating type to undergo sex.  As it is rare to encounter both mating types in nature, it is one of the reasons why it may be difficult to view the sexual cycle of some fungi.

The professor presented the form classes of Fungi Imperfecti: Blastomycetes, Hyphomycetes, Coelomycetes, and Mycelia Sterilia.  He showed various pictures of the form classes, many of which were in the form class Hyphomycetes, which displayed intricate mycelial conidial fungi.  The Coelomycetes were represented by some diverse conidiogenous multihyphal aggregates.

Finally, Dr. Szaniszlo closes his discussion of Fungi Imperfecti by listing some problems with the division Fungi Imperfecti.

(The professor moves on to Topic 3 for the remainder of Lecture Five)

End Lecture Five (2/3/2000)

Printed Notes, Main Points:

(click on the main point headings to jump to the selected location in the notes)

 

 


Review Definition of Fungi

Eukaryotic, heterotrophic and absorptive organisms, which have cell walls, typically reproduce asexually and/or sexually by producing spores, and grow either reproductively by budding or nonreproductively by hyphal tip elongation.

Definition excludes:

1. bacteria & blue-green algae (prokaryotes)

2. slime molds

3. true algae & higher plants

4. animals

5. etc.

Definition may include some non-fungi

1. Oomycetes (water molds)

2. etc.

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About Fungal Thalli (pl), Thallus (sing):

Many fungi characterized by a distinctive, multinucleate, vegetative (somatic) thallus (body) called

the mycelium* (singular)

mycelia* (plural).

The mycelium consists of a branching system of walled tubes called

hyphae (plural)

hypha (singular)

Mechanisms of hyphal growth** are

apical extension &

lateral branching.

* term usually used with filamentous fungi

**nonreproductive growth vs reproductive yeast growth

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Vegetative hyphal growth 
vs yeast growth

hyphal growth = apical extention

yeast growth= budding

hyphal growth= nonreproductive

yeast growth = reproductive

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Fungal nutrition

A. Absorptive mode

1. over whole surface or

2. via restricted absorbing regions , e.g.

a. rhizoids in "lower" fungi

b. substrate hyphae* in "higher" fungi

c. apical tips of hyphae

*The substrate hyphae of molds nourish the aerial hyphae and reproductive hyphae

B. Extracellular digestion

Fungi secrete enzymes that depolymerize complex natural products (proteins, carbohydrates, lipids, etc.) so they can be absorbed as sources of carbon and energy.

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Question: What are fungi?

1. Because of nutritional mode they are ecologically considered among the primary decomposers*.

2. Because of size, nutrition and history, they are considered to be microbes by microbiologists.

3. Because of structure and history, they have historically been considered to be plants by botanists.

Question: Are fungi plants?

No!…opinions of Whitaker, Margulis, Cavalier-Smith, Kendrick, PJS, etc.

* Reducers

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Questions

1. What was the pre-molecular biology basis for the reclassifications among the pre-Whittaker "fungi?"

2. What is the reality of the molecular revolution to the "new taxonomy" of the Kingdom Fungi?

2-1(new)

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Reasons Fungi not Plants:

1. Ultrastructural studies suggest uniqueness

- certainly more "primitive" fungi (Chytridiomycota) are not plants

- flagellation patterns of Chytridiomycota and other fungal-like protists suggest independent origins*

- derivation from nonphotosynthetic ancestors

* polyphyletic

2. Fungal organization is different from and nonhomologous with that of plants.

- convergent evolution -- similar structures and life cycles.

- tissue-like structure of hyphal origin*

* hyphal aggregates

3. Nutrition - fungi and most fungal-like protists were most-likely never photosynthetic.

Therefore, no more related to higher animals than higher plants.

Fungal Nutrition - Absorptive

Plant Nutrition - Photosynthetic

Animal Nutrition - Endocytotic or ingestive and absorptive

If not plants, then what?

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Kingdom Systems

Systems

Kingdoms

1. 2 kingdoms

1. Plantae

 

2. Animalatae

2. 3 kingdoms

1. Protista*

 

2. Metaphyta

 

3. Metazoa

3. 4 kingdoms

1. Monera

 

2. Protista*

 

3. Metaphyta

 

4. Metazoa

4. Whittaker's 5 kingdom system  (~ 1969)

 

*Concept #1 Protista = unicells or their colonial associations

*Concept #2 Protista = unicells or/and other organisms which lack tissue specialization of higher plants.

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What are fungi? - Question

Fungi are fungi. - Answer!

1) Fungi are accepted as the third kingdom of higher organisms whose origins flow naturally from a primitive eucaryotic protist.

2) Fungal origins are obscured but probably flowed from the Chytridiomycota, a group that we will study in this course, together with a few other fungal-like protists tradition ally studied by mycologists.

*Chytridiomycota are currently said to be most likely ancestor group

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Subdisciplines of taxonomy

1. Identification

Recognition of organism

2.* Classification

Recognition of relationships

(or nonrelationships)

3. Nomenclature

Correct naming of organisms (use of rules of botanical nomenclature)

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Classification hierarchy and fungal classification suffixes

Kingdom-Fungi

Subkingdom-mycotera

Division/phylum-mycota

Subdivision/subphylum-mycotina

Class-mycetes

Order-ales

Family-aceae

Genus, Saccharomyces

Species, S. cerevisiae

Organisms in the same taxon are more related than are organisms in different taxa.

17=2b

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Pre-Whitaker fungal classification (1969)

Kingdom - Plantae (Metaphyta)

Division - Eumycota

Class - Phycomycetes*

Class - Ascomycetes

Class - Basidiomycetes

Class - Deuteromycetes (Fungi Imperfecti)

* The so-called traditional "lower fungi"

*All produce their mitotically-derived reproductive cells in a cell called a sporangium. (sporangia)(pl)

* Lumping into single class erroneously suggested relatively close relationships

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Where are the Phycomycetes* today?

Kingdom - Protozoa**

Division - Plasmodiophoromycota

Kingdom - Chromista

Division - Hyphochytridiomycota

Division - Oomycota

Kingdom - Fungi

Subkingdom - Mastigomycotera***

Division - Chytridiomycota

Subkingdom - Amastigomycotera

Division - Zygomycota

*The lower fungi

**see lab manual, appendix a, for members, etc.

***Historically also:

Acrasiomycota - cellular slime molds

Labyrinthalomycota - gliding slime molds

Myxomycota - plasmodial slime molds, etc. (see other materials on reserve)

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Factors that originally led to the grouping of these different kinds of organisms into single class "Phycomycetes."

1. Their tendency to be vegetatively aseptate & multinucleate (coenocytic)

2. Their tendency to produce their mitotically derived, reproductive propagules (zoospores or sporangiospores) in a cell called a sporangium/sporangia (pl).

3. Their tendency to look alike and have superficially similar biologies that were clearly different than those of "higher fungi"

 

20 (revised 1/23/00)

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Major diagnostic characteristic of sporangial fungi and of some fungal-like protists

Phylum/Division

Sporangial propagule type

Plasmodiophoromycota

Zoospore* (1 whip-lash and one stub type rudiment of a flagellum

Hyphochytridiomycota

Zoospore (1 tinsel-type anterior flagellum)

Oomycota

Zoospore (1 tinsel- and 1 whip-lash flagellum)

Chytridiomycota

Zoospore (1 whip-lash posterior flagellum)

Zygomycota Sporangiospore**

 

* Zoospores are mycotically-derived reproductive cells lacking dormancy qualities and cell walls and having undulopodia.

** Sporangiospores have dormancy qualities and cell walls, but lack undulopodia

21=2-4 (rev. 1/23/00)

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Kingdom Protozoa

Division – Plasmodiophoromycota

Class - Plasmodiophoromycetes

Order - Plasmodiophorales**

* both hyphal and nonhyphal types

** intracellular plasmodial parasites of plants, algae and fungi - maybe more similar to animals than higher fungi

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Kingdom Chromista

Division - Hyphochytridiomycota*

Class - Hyphochytridiomycetes

Order - Hyphochytridiales

* Mostly parasites of algae and fungi or saprophytic plant and insect materials

* Nonhyphal and rudimentary hyphal representatives

Division - Oomycota**

Class - Oomycetes

Order - Saprolegniales

Order - Leptomitales

Order - Lagenidiales

Order - Olpidiopsidales

Order - Peronosporales

Order - Pythiales

** nonhyphal and hyphal members, orders in transition

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Kingdom Fungi

Kingdom - Fungi

Subkingdom - Mastigomycotera*

Division - Chytridiomycota**

Subkingdom - Amastigomycotera***

Division - Zygomycota

Subkingdom - Eumycotera

Division - Ascomycota

Division - Basidiomycota

Division - Fungi Imperfecti
(Deuteromycota)

*members produce zoospores

** presence of sporangia led to original inclusion among Phycomycetes

*** members produce sporangiospores

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Orders of Chytridiomycota

Kingdom - Fungi

Subkingdom - Mastigomycotera

Division - Chytridiomycota*

Class - Chytridiomycetes

Order - Chytridiales

Order - Spizellomycetales

Order - Blastocladiales

Order - Monoblephariadales

Order - Neocallimasticales

Order - Harpocyhytriales

* Both hyphal and nonhyphal types show signs of parallel evolutionary trends with Oocmycota members

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Classification of Zygomycota

Kingdom - Fungi

Subkingdom - Amastigomycotera*

Phylum - Zygomycota+

(cont below)

*Fungi that produce nonflagellate mitospores (sporangiospores**)  in a sporangium (species once classified among the phycomycetes)

**Sporangiospores (vs zoospores) have cell walls, dormancy qualities, no flagella, & tend to be wind disseminated

+Also characterized often by hyphae and the production of a karyospore (type of sexual spore) called a zygospore

(cont from above)

Class - Zygomycetes*

Order - Mucorales  (black bread molds)

Order - Entomophthorales (many insect pathogens)

Order - Zoopagales (insect gut symbionts)

Order - Endoganales  (endomycorrhizal fungi)

Order - Glomales (endomycorrhizal fungi)

Class – Trichomycetes**

 

* mostly hyphal organisms with a total of 7 orders

** 4 orders of poorly studied species that include numerous arthropod gut symbionts that are hyphal or rudimentarially hyphal. We will not study this group.

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Eumycotera Classification

Division

Major Characteristics(s)

Ascomycota

Ascus/Asci & Ascospores

Basidiomycota

Basidium/Basidia & Basidiospores**

Fungi Imperfecti

Absence of Sporangia, Asci or Basidia, Ascospores or Basidiospores

 

* ascospores = endogenous meiospores***

** basidiospores = exogenous meiospores***

*** true spores with cell walls and dormancy qualities.

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Eumycotera Classification (cont)

Kingdom Fungi

Subkingdom - Eumycotera*

Phylum - Ascomycota**

Phylum - Basidiomycota**

Phylum - Fungi Imperfecti***

* absence of mitosporangia (the higher fungi, the nonsporangial fungi)

** sometimes combined in one group (e.g. dikaryomycota).

*** a nonphylogenetic (artificial) phylum whose members may or may not be related (taxonomy does not reflect relationships)  The relationships among these fungi are being clarified by DNA and protein sequencing.

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Eumycotera Classification (cont)

Division

Major Characteristics(s)

Ascomycota

Meiosporangium called an ascus, which produces endo-genous meiospores called ascospores

Basidiomycota

Meiosporangium called a basidium, which produces exo-genous meiospores called basidiospores*

Fungi Imperfecti

No known or observed meiosporangia, therefore no ascospores or basidio-spores, therefore no absolute ability to classify morphologically.**

*true spores with dormancy qualities
**possibly can now by molecular biology

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Ascomycota classification

Division - Ascomycota

Subdivision - Hemiascomycotina*

Class - Hemiascomycetes

Order - Saccharomycetales
(Ascus from diploid nucleus w/o N+N state)

Class - Archaeascomycetes

Order - Taphrinales
(Ascus from N+N cell)

Order - Schizosaccharomycetales  (fission yeasts)

Order - Pneumocystidales (for Pneumocystis carinii, etc.)

Order - Protomycetales

*lack ascocarps (multihyphal tissue-like structures surrounding their asci. both hyphal & nonhyphal members.

Subdivision - Euascomycotina**

**have ascocarps associated with their asci.

N+N = dikaryon state                                                                                                         46

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Classes of Euascomycotina

Class

Ascoma or Ascocarp type

1. Plectomycetes

Cleistothecia (a)

2. Pyrenomycetes

Perithecia (b)

3. Discomycetes

Apothecia (c)

4. Loculoascomycetes

Ascostroma (d)

5. Laboulbeniomycetes

Very specialized Perithecia

 

a. closed ascocarps
b. flask-shaped ascocarps
c. cup or goblet shaped ascocarps
d. stroma with locuoles (cavities)

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Basidiomycota Classification

Subkingdom - Eumycotera

Phylum - Basidiomycota

Subphylum - Heterobasidiomycotina*,**

Class - Urediniomycetes (rusts)

Order - Uredinales

Class - Ustomycetes (smuts)

Order - Ustilaginales

* basidia from teliospores (dikaryotic spores)
**no  "mushroom-like" basidiocarps

Subphylum - Holobasidiomycotina

Class - Phragmobasidiomycetes* (5 orders)

Order - Tremellales
(have cruciately septate Basidia)

Order - Auriculariales
(have transversely septate Basidia)

* have septate basidia

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Basidiomycota Classification (cont.)

Subphylum - Holobasidiomycotina * (cont.)

Class – Holobasidiomycetes (27 orders)

Order - Dacrymycetales
(have tuning-fork type basidium)

Order - Tulasnellales (have Holobasidia with swollen sterigmata)

Order - Agaricales (the mushrooms)

Order – Boletales (the fleshy pore fungi)

Order – Cantharelles (chantarelles, and tooth fungi, etc.)

Order - Exobasidials

Order - Gautieriales

Order - Hymenogastrales

Order - Lycoperdales (the puffballs, earth stars, etc.)

Order - Melanogastrales

Order - Nidulariales (the bird's nest fungi)

Order - Phallales (the stinkhorns)

Order - Porales (woody shelf fungi or woody bracket fungi)

Order – Thelephorales (the coral and leather fungi)

Order – Sclerodermatales (the earth balls)

 

* most have typical holobasidum, but different kinds of basidiocarps

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Review - Kingdom Fungi

Kingdom – Fungi

Subkingdom - Mastigomycotera

Phylum - Chytridiomycota

Subkingdom - Amastigomycotera

Phylum - Zygomycota

Class - Zygomycetes

Orders- Mucorales (etc)

Class - Trichomycetes

Orders - 1 to 4

Subkingdom - Eumycotera

Phylum - Ascomycota

Subphylum - Hemiascomycotina

Class - Hemiascomycetes

Order - Saccharomycetales
(Ascus from diploid nucleus w/o N+N state)

Class - Archaeascomycetes

Order - Taphrinales
(Ascus from N+N cell)

Order - Schizosaccharomycetales  (fission yeasts)

Order - Pneumocystidales (for Pneumocystis carionii, etc.)

Order – Protomycetales

Subphylum - Euascomycotina

Class - Plectomycetes

Class - Pyrenomycetes

Class - Discomycetes

Class - Loculoascomycetes

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Review - Kingdom Fungi (cont.)

Subkingdom - Eumycotera

Phylum - Basidiomycota

Subphylum - Heterobasidiomycotina

Class - Urediniomycetes

Order - Uredinales

Class - Ustomycetes

Order - Ustilaginales

Subphylum - Heterobasidiomycotina

Class - Phragomobasidiomycetes

Orders - Tremelales*, Auriculariales

Class - Holobasidiomyetes

Orders -  Dacrymycetales, Tulasnellales, Exobasideals, Aphyllophorales, Hymenogastrales, Gautieriales, Phallales, Lycoperdales, Tulostomatales, Sclerodermatatales, Nidulareales, Agaricales, Melanogadstrales, Porales

See my reference guide to Fungi & Fungal-like Protists (circa 1999), & handout
*Current most accepted order for F.(C) neoformans but may be a member of the Ustomycetes class

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FUNGI IMPERFECTI
(Deuteromycota)

Nonphylogenetic group of fungi without known sexual cycles or with unobserved sexual cycles.

Unique Phylum/Division

1.  Groupings may or may not represent phylogeny e.g., species in same genus may be less related than species in different genera

2.  Most members probably should be included in another Division, particularly Ascomycota and Basidiomycota

3.  Members often have two and sometimes more acceptable scientific names

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WHY FUNGI IMPERFECTI?

1.  Recognition that sexual cycles of fungi, particularly of Ascomycota and Basidiomycota, were important for understanding relationships created the problem

2.  Problem was what to do taxonomically with fungi that had no known sex cycles?*

3.  Problem solved by creating a Division of fungi in which fungi are named with little regard for relationships.  Identification without regard to classification; based on observations and naming of asexual states.

*majority

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Question - Why are asexual states usually known before sexual states?

1.  If organism grows then it usually exhibits its asexual (anamorphic) phase first.

asexual (anamorphic) phase first

vegetative growth ------> asexual

reproduction -------------> sexual reproduction

2.  With practically important fungi, one can identitfy and work with fungus without knowing about sexual stage.

3.  Induction of sexuality is often dependent upon inducing sex in mated strains.

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Homothallism vs Heterothallism

Many fungi are heterothallic*

Haploid self-sterile (nonsexual) strains
                                                                              A***                A A ****
A ( haploid/1N)                                     (1N)        A                     A A
X--------------------> Aa --------> meiosis** -------> mitosis ----->
a ( haploid/1N)     (2N)                                         a                     a a
                                                                              a                     a a

Only a few are homothallic (self-fertile)

*     sex requires = pairs of opposite mating strains
**   meiosis often requires special physiological conditions
*** meitotic nuclei incorporated into
meiospores
ascospores
basidiospores

**** somtimes, etc.

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ASEXUAL STATES = IMPERFECT STATE

Therefore:  Fungi Imperfecti = Deuteromycota

1.  Fungi Imperfecti named according to rules of Botanical Nomenclature for asexual fungi

2.  When perfect states (sexual) become known they are classified and renamed according to rules of Botanical Nomenclature for sexual fungi

Older Fungi Imperfecti names usually better known

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Review Fungi Imperfecti 

1. Fungi having no known or observed sexual state (teleomorph)

2. Usually asexual states (anamorphs) of ascomycota or basidiomycota

3. When sexual state of an imperfect (asexual) fungus is observed and described the fungus is

a. renamed according to rules of botanical nomenclature

b. reclassified according to characteristics of the holomorph = whole fungus considering biology of both anamorph and teleomorph

4. Until discovery of teleomorph fungus is a member of the fungi imperfecti and included in its "hierarchy" of categorization.

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FUNGI IMPERFECTI 

Hierarchy of Categorization* 

Form - Class

Form - Order

Form - Families

Form - Genera

Form - Species

*System useful for communication and identification, but does not necessarily reflect relationships (phylogeny)

*Not a classification system

1. Form class - Blastomycetes

nonsexual yeasts

2. Form class - Hyphomycetes

Mycelial (hyphal) conidial fungi

3. Form class - Coelomycetes

Mycelial (hyphal) conidial fungi with conidiogenous structures associated with multihyphal aggregates.**

4. Form Class - Mycelia sterilia 

nonsporulating hyphal fungi.

**  Conidioma

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Examples of common anamorphic genera (Fungi imperfecti) whose teleomorphic names are often established but seldom used.

Anamorphic name

Teleomorphic name

Penicillium

Telaromyces, etc.

Aspergillus

Eurotium, etc.

Rhodotorula

Rhodosporidium

Trichophyton

Arthroderma

Microsporum

Arthroderma

Blastomyces

Ajellomyces

Cryptococcus

Filobasidiella

Candida

Saccharomyces

Pichia

Hansenula etc

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Today's Problems with Division Fungi Imperfecti

Good News - perfect states of many fungi have been, and continue to be, discovered

Bad News - hundreds of common fungi have new names

More Bad News - great resistance to learning and understanding about why we are having reclassifications*

Great resistance to use of new names by non-mycologists and professionals.

Both imperfect and perfect names of sexual fungi commonly used.

Particularly true in regard to applied fields like

·        medical mycology

·        plant pathology

·        industrial mycology

* will be compounded by molecular classifications

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