c

A S
  (pronounced ~ S
~Ú is a eukaryotic organism that is a member of the kingdom

©
 (pronounced ~ S   ~Ú
 The fungi are heterotrophic organisms possessing a
chitinous cell wall The majority of species grow as multicellular filaments called hyphae
forming a mycelium; some fungal species also grow as single cells Sexual and asexual
reproduction of the fungi is commonly via spores, often produced on specialized structures or in
fruiting bodies Some species have lost the ability to form specialized reproductive structures,
and propagate solely by vegetative growth Yeasts, molds, and mushrooms are examples of
fungi The fungi are a monophyletic group that is phylogenetically clearly distinct from the
morphologically similar slime molds (myxomycetesÚ and water molds (oomycetesÚ The fungi
are more closely related to animals than plants, yet the discipline of biology devoted to the study
of fungi, known as mycology, often falls under a branch of botany

Occurring worldwide, most fungi are largely invisible to the naked eye, living for the most part
in soil, dead matter, and as symbionts of plants, animals, or other fungi They perform an
essential role in all ecosystems in decomposing organic matter and are indispensable in nutrient
cycling and exchange Some fungi become noticeable when fruiting, either as mushrooms or
molds Many fungal species have long been used as a direct source of food, such as mushrooms
and truffles and in fermentation of various food products, such as wine, beer, and soy sauce
More recently, fungi are being used as sources for antibiotics used in medicine and various
enzymes, such as cellulases, pectinases, and proteases, important for industrial use or as active
ingredients of detergents Many fungi produce bioactive compounds called mycotoxins, such as
alkaloids and polyketides that are toxic to animals including humans Some fungi are used
recreationally or in traditional ceremonies as a source of psychotropic compounds Several
species of the fungi are significant pathogens of humans and other animals, and losses due to
diseases of crops (e g , rice blast diseaseÚ or food spoilage caused by fungi can have a large
impact on human food supply and local economies

G



hide

6c u Etymology and definition

6c  Diversity
6c º Importance for human use
c º u Cultured foods
c º  Other human uses
c º º Mycotoxins
c º  Edible and poisonous fungi
c º  Fungi in the biological control of pests
c º  Bioremediation
6c  Ecology
c  u Symbiosis
rc  u u With plants
rc  u  With insects
rc  u º As pathogens and parasites
c   Nutrition and possible autotrophy
6c  Morphology
c  u Microscopic structures
c   Macroscopic structures
c  º Morphological and physiological features for substrate penetration
6c  Reproduction
c  u Asexual reproduction
c   Sexual reproduction
c  º Spore dispersal
c   Other sexual processes
6c  Phylogeny and classification
c  u Physiological and morphological traits
c   Evolutionary history
rc   u Cladogram
c  º The taxonomic groups of fungi
c   Phylogenetic relationships with other fungus-like organisms
6c m See also
6c  Notes and references
6c u Further reading
6c uu External links

Œ


S


The English word S
  is directly adopted from the Latin S
 , meaning "mushroom", used
in Horace and Pliny
º This in turn is derived from the Greek word 
/ıijȠȖȖȠȢ
("sponge"Ú, referring to the macroscopic structures and morphology of some mushrooms and
molds and also used in other languages (e g , the German 
 ("sponge"Ú or 
 
for some types of mushroomÚ
£ 
Fungi have a worldwide distribution, and grow in a wide range of habitats, including deserts
Most fungi grow in terrestrial environments, but several species occur only in aquatic habitats
Fungi along with bacteria are the primary decomposers of organic matter in most if not all
terrestrial ecosystems worldwide Based on observations of the ratio of the number of fungal
species to the number of plant species in some environments, the fungal kingdom has been
estimated to contain about u  million species
 Around  , fungal species have been
formally described by taxonomists, but the true dimension of fungal diversity is still unknown

Most fungi grow as thread-like filaments called hyphae, which form a mycelium, while others
grow as single cells

 Until recently many fungal species were described based mainly on
morphological characteristics, such as the size and shape of spores or fruiting structures, and
biological species concepts; the application of molecular tools, such as DNA sequencing, to
study fungal diversity has greatly enhanced the resolution and added robustness to estimates of
diversity within various taxonomic groups
m

  

S 

 

Sacharomyces cerevisiae cells in DIC microscopy

Human use of fungi for food preparation or preservation and other purposes is extensive and has
a long history: yeasts are required for fermentation of beer, wine
 and bread, some other fungal
species are used in the production of soy sauce and tempeh Mushroom farming and mushroom
gathering are large industries in many countries Many fungi are producers of antibiotics,
including ȕ-lactam antibiotics such as penicillin and cephalosporin
u  Widespread use of these
antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, and
many others began in the early  th century and continues to play a major part in anti-bacterial
chemotherapy The study of the historical uses and sociological impact of fungi is known as
ethnomycology

G
 S 

Baker's yeast or 

 
, a single-cell fungus, is used in the baking of bread
and other wheat-based products, such as pizza and dumplings
uu Several yeast species of the
genus Saccharomyces are also used in the production of alcoholic beverages through
fermentation
u Mycelial fungi, such as the shoyu koji mold (   
Ú, are used in the
brewing of Shoyu (soy sauceÚ and preparation of tempeh
uº Quorn is a high-protein product
made from the mold, © 
 


 , and is used in vegetarian cooking

 

 

Fungi are also used extensively to produce industrial chemicals like lactic acid, antibiotics and
even to make stonewashed jeans
u Several fungal species are ingested for their psychedelic
properties, both recreationally and religiously (see main article, ]
  Ú

0  


0


0


Many fungi produce compounds with biological activity Several of these compounds are toxic
and are therefore called mycotoxins, referring to their fungal origin and toxic activity Of
particular relevance to humans are those mycotoxins that are produced by moulds causing food
spoilage and poisonous mushrooms (see belowÚ Particularly infamous are the aflatoxins, which
are insidious liver toxins and highly carcinogenic metabolites produced by    species
often growing in or on grains and nuts consumed by humans, and the lethal amatoxins produced
by mushrooms of the genus 


Other notable mycotoxins include ochratoxins, patulin,
ergot alkaloids, and trichothecenes and fumonisins, all of which have significant impact on
human food supplies or animal livestock
u

Mycotoxins belong to the group of secondary metabolites (or natural productsÚ Originally, this
group of compounds had been thought to be mere byproducts of primary metabolism, hence the
name "secondary" metabolites However, recent research has shown the existence of biochemical
pathways solely for the purpose of producing mycotoxins and other natural products in fungi
u
Mycotoxins provide a number of fitness benefits to the fungi that produce them in terms of
physiological adaptation, competition with other microbes and fungi, and protection from
fungivory
u
um These fitness benefits and the existence of dedicated biosynthetic pathways for
mycotoxin production suggest that the mycotoxins are important for fungal persistence and
survival

Œ


 
S


Asian mushrooms, clockwise from left, enokitake, buna-shimeji, bunapi-shimeji, king oyster
mushroom and shiitake
Black Périgord Truffle (  

 Ú, cut in half

Stilton cheese veined with ]
   S 

Some of the best known types of fungi are the edible and the poisonous mushrooms Many
species are commercially raised, but others must be harvested from the wild 
  ,
sold as button mushrooms when small or Portobello mushrooms when larger, are the most
commonly eaten species, used in salads, soups, and many other dishes Many Asian fungi are
commercially grown and have gained in popularity in the West They are often available fresh in
grocery stores and markets, including straw mushrooms (



Ú, oyster
mushrooms (]   
 Ú, shiitakes (


Ú, and enokitake (©
 

spp Ú

There are many more mushroom species that are harvested from the wild for personal
consumption or commercial sale Milk mushrooms, morels, chanterelles, truffles, black trumpets,
and  
 mushrooms (  Ú (also known as king boletesÚ all demand a high price on
the market They are often used in gourmet dishes

For certain types of cheeses, it is also a common practice to inoculate milk curds with fungal
spores to foment the growth of specific species of mold that impart a unique flavor and texture to
the cheese This accounts for the blue colour in cheeses such as Stilton or Roquefort which is
created using ]
   S  spores
u Molds used in cheese production are usually non-
toxic and are thus safe for human consumption; however, mycotoxins (e g , aflatoxins,
roquefortine C, patulin, or othersÚ may accumulate due to fungal spoilage during cheese ripening
or storage
 

Many mushroom species are toxic to humans, with toxicities ranging from slight digestive
problems or allergic reactions as well as hallucinations to severe organ failures and death Some
of the most deadly mushrooms belong to the genera 
,  

 , and most infamously,



The latter genus includes the destroying angel ! 
" and the death cap !

", the most common cause of deadly mushroom poisoning
u The false morel
(# 
 

Ú is considered a delicacy by some when cooked, yet can be highly toxic
when eaten raw
  
   was considered edible until being implicated in some
serious poisonings causing rhabdomyolysis
º

Fly agaric mushrooms (! 


Ú also cause occasional poisonings, mostly as a result of
ingestion for use as a recreational drug for its hallucinogenic properties Historically Fly agaric
was used by Celtic Druids in Northern Europe and the Koryak people of north-eastern Siberia for
religious or shamanic purposes
 It is difficult to identify a safe mushroom without proper
training and knowledge, thus it is often advised to assume that a mushroom in the wild is
poisonous and not to consume it

©






 S

In agricultural settings, fungi that actively compete for nutrients and space with, and eventually
prevail over, pathogenic microorganisms, such as bacteria or other fungi, via the competitive
exclusion principle,
 or are parasites of these pathogens, may be beneficial agents for human
use For example, some fungi may be used to suppress growth or eliminate harmful plant
pathogens, such as insects, mites, weeds, nematodes and other fungi that cause diseases of
important crop plants
 This has generated strong interest in the use and practical application of
these fungi for the biological control of these agricultural pests Entomopathogenic fungi can be
used as biopesticides, as they actively kill insects
 Examples of fungi that have been used as
biological insecticides are 
 




, 0
 


, $  
spp,
]
 spp, and   


m
 Endophytic fungi of grasses of the genus
% , such as %!

  produce alkaloids that are toxic to a range of
invertebrate and vertebrate herbivores These alkaloids protect the infected grass plants from
herbivory, but some endophyte alkaloids can cause poisoning of grazing animals, such as cattle
and sheep
º  Infection of grass cultivars of turf or forage grasses with isolates of the grass
endophytes that produce only specific alkaloids to improve grass hardiness and resistance to
herbivores such as insects, while being non-toxic to livestock, is being used in grass breeding
programs
ºu

D  



0


0 


Certain fungi, in particular 'white rot' fungi, can degrade insecticides, herbicides,
pentachlorophenol, creosote, coal tars, and heavy fuels and turn them into carbon dioxide, water,
and basic elements
º Research has recently discovered that fungi can be used to lock uranium
into mineral form
ºº

Œ

Polypores growing on a tree in Borneo

Although often inconspicuous, fungi occur in every environment on Earth and play very
important roles in most ecosystems Along with bacteria, fungi are the major decomposers in
most terrestrial (and some aquaticÚ ecosystems, and therefore play a critical role in
biogeochemical cycles and in many food webs As decomposers, they play an indispensable role
in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to
inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other
organisms
º
º

  

Many fungi have important symbiotic relationships with organisms from most if not all
Kingdoms
º
º
ºm These interactions can be mutualistic or antagonistic in nature, or in case of
commensal fungi are of no apparent benefit or detriment to the host
º
 
u






Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant-fungus
associations and is of significant importance for plant growth and persistence in many
ecosystems; over 
of all plant species engage in some kind of mycorrhizal relationship with
fungi and are dependent upon this relationship for survival

º
 The mycorrhizal symbiosis
is ancient, dating to at least  million years ago
 It often increases the plant's uptake of
inorganic compounds, such as nitrate and phosphate from soils having low concentrations of
these key plant nutrients
º In some mycorrhizal associations, the fungal partners may mediate
plant-to-plant transfer of carbohydrates and other nutrients Such mycorrhizal communities are
called "common mycorrhizal networks"


Lichens are formed by a symbiotic relationship between algae or cyanobacteria (referred to in

lichens as "photobionts"Ú and fungi (mostly various species of ascomycetes and a few
basidiomycetesÚ, in which individual photobiont cells are embedded in a tissue formed by the
fungus
 As in mycorrhizas, the photobiont provides sugars and other carbohydrates, while the
fungus provides minerals and water The functions of both symbiotic organisms are so closely
intertwined that they function almost as a single organism



Many insects also engage in mutualistic relationships with various types of fungi Several groups
of ants cultivate fungi in the order Agaricales as their primary food source, while ambrosia
beetles cultivate various species of fungi in the bark of trees that they infest
m Termites on the
African Savannah are also known to cultivate fungi



 






However, many fungi are parasites on plants, animals (including humansÚ, and other fungi
Serious fungal pathogens of many cultivated plants causing extensive damage and losses to
agriculture and forestry include the rice blast fungus 0


  
,
  tree pathogens
such as &
  and &

'  causing Dutch elm disease,
u and
 
 



responsible for chestnut blight,
 and plant-pathogenic fungi in the
genera © 
 , (
, 


, and  ;
  fungi with the potential to cause
serious human diseases, especially in persons with immuno-deficiencies, are in the genera
  , 


,   ,
º
u
 $

,
 and ]
 
 Several
pathogenic fungi are also responsible for relatively minor human diseases, such as athlete¶s foot
and ringworm Some fungi are predators of nematodes, which they capture using an array of
specialized structures, such as constricting rings or adhesive nets


   


 

  

Growth of fungi as hyphae on or in solid substrates or single cells in aquatic environments is

adapted to efficient extraction of nutrients from these environments, because these growth forms
have high surface area to volume ratios These adaptations in morphology are complemented by
hydrolytic enzymes secreted into the environment for digestion of large organic molecules, such
as polysaccharides, proteins, lipids, and other organic substrates into smaller molecules
m

 
These molecules are then absorbed as nutrients into the fungal cells

Traditionally, the fungi are considered heterotrophs, organisms that rely solely on carbon fixed
by other organisms for metabolism Fungi have evolved a remarkable metabolic versatility that
allows many of them to use a large variety of organic substrates for growth, including simple
compounds as nitrate, ammonia, acetate, or ethanol
u
 Recent research raises the possibility
that some fungi utilize the pigment melanin to extract energy from ionizing radiation, such as
gamma radiation for "radiotrophic" growth
º It has been proposed that this process might bear
some similarity to photosynthesis in plants,
º but detailed biochemical data supporting the
existence of this hypothetical pathway are presently lacking

0 

0    
Mold covering a decaying peach over a period of six days The frames were taken approximately
u hours apart

Though fungi are part of the opisthokont clade, all phyla except for the chytrids have lost their
posterior flagella
 Fungi are unusual among the eukaryotes in having a cell wall that, besides
glucans (e g , ȕ-u,º-glucanÚ and other typical components, contains the biopolymer chitin


Many fungi grow as thread-like filamentous microscopic structures called hyphae, and an
assemblage of intertwined and interconnected hyphae is called a mycelium
 Hyphae can be
septate, i e , divided into hyphal compartments separated by a  , each compartment
containing one or more nuclei or can be coenocytic, i e , lacking hyphal compartmentalization
However, septa have pores, such as the doliporus in the basidiomycetes that allow cytoplasm,
organelles, and sometimes nuclei to pass through
 Coenocytic hyphae are essentially
multinucleate supercells
 In some cases, fungi have developed specialized structures for
nutrient uptake from living hosts; examples include haustoria in plant-parasitic fungi of nearly all
divisions, and arbuscules of several mycorrhizal fungi,
 which penetrate into the host cells for
nutrient uptake by the fungus

0
    

Fungal mycelia can become visible macroscopically, for example, as concentric rings on various
surfaces, such as damp walls, and on other substrates, such as spoilt food (see figureÚ, and are
commonly and generically called   (American spelling, Ú; fungal mycelia grown on
solid agar media in laboratory petri dishes are usually referred to as colonies, with many species
exhibiting characteristic macroscopic growth morphologies and colours, due to spores or
pigmentation

Specialized fungal structures important in sexual reproduction are the apothecia, perithecia, and
cleistothecia in the ascomycetes, and the fruiting bodies of the basidiomycetes, and a few
ascomycetes These reproductive structures can sometimes grow very large, and are well known
as mushrooms

0 







S
 S  





Fungal hyphae are specifically adapted to growth on solid surfaces and within substrates, and can
exert astoundingly large penetrative mechanical forces The plant pathogen, 0


 
 
, forms a structure called an appressorium specifically designed for penetration of plant
tissues, and the pressure generated by the appressorium, which is directed against the plant
epidermis can exceed m MPa (m barsÚ
m The generation of these mechanical pressures is the
result of an interplay between physiological processes to increase intracellular turgor by
production of osmolytes such as glycerol, and the morphology of the appressorium


·  


Fungi on a fence post near Orosí, Costa Rica

Reproduction of fungi is complex, reflecting the heterogeneity in lifestyles and genetic make up
within this group of organisms
 Many fungi reproduce either sexually or asexually, depending
on conditions in the environment These conditions trigger genetically determined developmental
programs leading to the expression of specialized structures for sexual or asexual reproduction
These structures aid both reproduction and efficient dissemination of spores or spore-containing
propagules



   


Asexual reproduction via vegetative spores or through mycelial fragmentation is common in

many fungal species and allows more rapid dispersal than sexual reproduction In the case of the
"Fungi imperfecti" or Deuteromycota, which lack a sexual cycle, it is the only means of
propagation Asexual spores, upon germination, may found a population that is clonal to the
population from which the spore originated, and thus colonize new environments



   


Sexual reproduction with meiosis exists in all fungal phyla, except the Deuteromycota It differs
in many aspects from sexual reproduction in animals or plants Many differences also exist
between fungal groups and have been used to discriminate fungal clades and species based on
morphological differences in sexual structures and reproductive strategies Experimental crosses
between fungal isolates can also be used to identify species based on biological species concepts
The major fungal clades have initially been delineated based on the morphology of their sexual
structures and spores; for example, the spore-containing structures, asci and basidia, can be used
in the identification of ascomycetes and basidiomycetes, respectively Many fungal species have
elaborate vegetative incompatibility systems that allow mating only between individuals of
opposite mating type, while others can mate and sexually reproduce with any other individual or
itself Species of the former mating system are called heterothallic, and of the latter homothallic

 

Most fungi have both a haploid and diploid stage in their life cycles In all sexually reproducing
fungi, compatible individuals combine by cell fusion of vegetative hyphae by anastomosis,
required for the initiation of the sexual cycle Ascomycetes and basidiomycetes go through a
dikaryotic stage, in which the nuclei inherited from the two parents do not fuse immediately after
cell fusion, but remain separate in the hyphal cells (see heterokaryosisÚ

In ascomycetes, dikaryotic hyphae of the hymenium form a characteristic ) at the hyphal
septum During cell division formation of the hook ensures proper distribution of the newly
divided nuclei into the apical and basal hyphal compartments An ascus (plural
Ú is then
formed, in which karyogamy (nuclear fusionÚ occurs These asci are embedded in an ascocarp, or
fruiting body, of the fungus Karyogamy in the asci is followed immediately by meiosis and the
production of ascospores The ascospores are disseminated and germinate and may form a new
haploid mycelium
u

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes Compatible haploid

hyphae fuse to produce a dikaryotic mycelium However, the dikaryotic phase is more extensive
in the basidiomycetes, in many cases also present in the vegetatively growing mycelium A
specialized anatomical structure, called a 



, is formed at each hyphal septum As
with the structurally similar hook in the ascomycetes, formation of the clamp connection in the
basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the
dikaryotic stage with two genetically different nuclei in each hyphal compartment
u A
basidiocarp is formed in which club-like structures known as basidia generate haploid
basidiospores after karyogamy and meiosis
 The most commonly known basidiocarps are
mushrooms, but they may also take many other forms (see Morphology sectionÚ

In zygomycetes, haploid hyphae of two individuals fuse, forming a zygote, which develops into a
zygospore When the zygospore germinates, it quickly undergoes meiosis, generating new
haploid hyphae, which in turn may form asexual sporangiospores These sporangiospores are
means of rapid dispersal of the fungus and germinate into new genetically identical haploid
fungal colonies, able to mate and undergo another sexual cycle followed by the generation of
new zygospores, thus completing the lifecycle

  



Both asexual and sexual spores or sporangiospores of many fungal species are actively dispersed
by forcible ejection from their reproductive structures This ejection ensures exit of the spores
from the reproductive structures as well as travelling through the air over long distances Many
fungi thereby possess specialized mechanical and physiological mechanisms as well as spore-
surface structures, such as hydrophobins, for spore ejection These mechanisms include, for
example, forcible discharge of ascospores enabled by the structure of the ascus and accumulation
of osmolytes in the fluids of the ascus that lead to explosive discharge of the ascospores into the
air
º The forcible discharge of single spores termed 
  involves formation of a
small drop of water (Buller's dropÚ, which upon contact with the spore leads to its projectile
release with an initial acceleration of more than u , g
 Other fungi rely on alternative
mechanisms for spore release, such as external mechanical forces, exemplified by puffballs
Attracting insects, such as flies, to fruiting structures, by virtue of their having lively colours and
a putrid odour, for dispersal of fungal spores is yet another strategy, most prominently used by
the stinkhorns

 

 

Besides regular sexual reproduction with meiosis, some fungal species may exchange genetic
material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of
fungal cells The frequency and relative importance of parasexual events is unclear and may be
lower than other sexual processes However, it is known to play a role in intraspecific
hybridization
 and is also likely required for hybridization between fungal species, which has
been associated with major events in fungal evolution








S



The mushroom & 




 eats wood

For a long time taxonomists considered fungi to be members of the Plant Kingdom This early
classification was based mainly on similarities in lifestyle: both fungi and plant are mainly
sessile, have similarities in general morphology and growth habitat (like plants, fungi often grow
in soil, in the case of mushrooms forming conspicuous fruiting bodies, which sometimes bear
resemblance to plants such as mossesÚ Moreover, both groups possess a cell wall, which is
absent in the Animal Kingdom However, the fungi are now considered a separate kingdom,
distinct from both plants and animals, from which they appear to have diverged approximately
one billion years ago
 Many studies have identified several distinct morphological,
biochemical, and genetic features in the Fungi, clearly delineating this group from the other
kingdoms For these reasons, the fungi are placed in their own kingdom






 





Similar to animals and unlike most plants, fungi lack the capacity to synthesize organic carbon
by chlorophyll-based photosynthesis; whereas plants store the reduced carbon as starch, fungi,
like animals and some bacteria, use glycogen
m for storage of carbohydrates A major
component of the cell wall in many fungal species is the nitrogen-containing carbohydrate,
chitin,
 also present in some animals, such as the insects and crustaceans, while the plant cell
wall consists chiefly of the carbohydrate cellulose The defining and unique characteristics of
fungal cells include growth as hyphae, which are microscopic filaments of between -u microns
in diameter and up to several centimetres in length, and which combined form the fungal
mycelium Some fungi, such as yeasts, grow as single ovoid cells, similar to unicellular algae
and the protists

Unlike many plants, most fungi lack an efficient vascular system, such as xylem or phloem for
long-distance transport of water and nutrients; as an example for convergent evolution, some
fungi, such as  

, form rhizomorphs or mycelial cords,
m  resembling and functionally
related to, but morphologically distinct from, plant roots

Some characteristics shared between plants and fungi include the presence of vacuoles in the
cell,
mu and a similar pathway in the biosynthesis of terpenes using mevalonic acid and
pyrophosphate as biochemical precursors; plants however use an additional terpene biosynthesis
pathway in the chloroplasts that is apparently absent in fungi
m Ancestral traits shared among
members of the fungi include chitinous cell walls and heterotrophy by absorption
u A further
characteristic of the fungi that is absent from other eukaryotes, and shared only with some
bacteria, is the biosynthesis of the amino acid, L-lysine, via the Į-aminoadipate pathway
mº

Similar to plants, fungi produce a plethora of secondary metabolites functioning as defensive

compounds or for niche adaptation; however, biochemical pathways for the synthesis of similar
or even identical compounds often differ markedly between fungi and plants
m
m

Œ


 

0


* 
SS


The first organisms having features typical of fungi date to u, million years ago, the
Proterozoic
m However, fungal fossils do not become common and uncontroversial until the
early Devonian, when they are abundant in the Rhynie chert
m

Even though traditionally included in many botany curricula and textbooks, fungi are now
thought to be more closely related to animals than to plants and are placed with the animals in
the monophyletic group of opisthokonts
u For much of the Paleozoic Era, the fungi appear to
have been aquatic, and consisted of organisms similar to the extant Chytrids in having flagellum-
bearing spores
mm The early fossil record of the fungi is fragmentary, to say the least The fungi
probably colonized the land during the Cambrian, long before land plants
m All modern classes
of fungi were present in the Late Carboniferous (Pennsylvanian EpochÚ
m For some time after
the Permian-Triassic extinction event, a fungal spike, originally thought to be an extraordinary
abundance of fungal spores in sediments formed shortly after this event, suggested that they were
the dominant life form during this period²nearly u
of the fossil record available from this
period
  However, the relative proportion of fungal spores relative to spores formed by algal
species is difficult to assess,
u the spike did not appear world-wide,

º and in many places it
did not fall on the Permian-Triassic boundary


Analyses using molecular phylogenetics support a monophyletic origin of the Fungi
m The
taxonomy of the Fungi is in a state of constant flux, especially due to recent research based on
DNA comparisons These current phylogenetic analyses often overturn classifications based on
older and sometimes less discriminative methods based on morphological features and biological
species concepts obtained from experimental matings



There is no unique generally accepted system at the higher taxonomic levels and there are
constant name changes at every level, from species upwards However, efforts among fungal
researchers are now underway to establish and encourage usage of a unified and more consistent
nomenclature
m Fungal species can also have multiple scientific names depending on its life
cycle and mode (sexual or asexualÚ of reproduction Web sites such as Index Fungorum and ITIS
define preferred up-to-date names (with cross-references to older synonymsÚ, but do not always
agree with each other

G

 


(
)



&)









©
  





%



+

# 

,)








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The major divisions (phylaÚ of fungi have been classified based mainly on their sexual
reproductive structures Currently, seven fungal divisions are proposed:
m

   
 
seen under microscope Flax root cortical cells containing paired
arbuscules
Conidiophores of molds of the genus   , an ascomycete, seen under microscope

6c The Chytridiomycota are commonly known as chytrids These fungi are ubiquitous with
a worldwide distribution; chytrids produce zoospores that are capable of active
movement through aqueous phases with a single flagellum Consequently, some
taxonomists had earlier classified them as protists on the basis of the flagellum
Molecular phylogenies, inferred from the rRNA-operon sequences representing the umS,
mS, and  mS ribosomal subunits, suggest that the Chytrids are a basal fungal group
divergent from the other fungal divisions, consisting of four major clades with some
evidence for paraphyly or possibly polyphyly
mm
6c The Blastocladiomycota were previously considered a taxonomic clade within the
Chytridiomycota Recent molecular data and ultrastructural characteristics, however,
place the Blastocladiomycota as a sister clade to the Zygomycota, Glomeromycota, and
Dikarya (Ascomycota and BasiomycotaÚ The blastocladiomycetes are fungi that are
saprotrophs and parasites of all eukaryotic groups and undergo sporic meiosis unlike their
close relatives, the chytrids, which mostly exhibit zygotic meiosis
mm
6c The Neocallimastigomycota were earlier placed in the phylum Chytridomycota Members
of this small phylum are anaerobic organisms, living in the digestive system of larger
herbivorous mammals and possibly in other terrestrial and aquatic environments They
lack mitochondria but contain hydrogenosomes of mitochondrial origin As the related
chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or
polyflagellate
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6c The Zygomycota contain the taxa, Zygomycetes and Trichomycetes, and reproduce
sexually with meiospores called zygospores and asexually with sporangiospores Black
bread mold (· 
S Ú is a common species that belongs to this group; another
is ] , which is capable of ejecting spores several meters through the air Medically
relevant genera include 0  , ·  , and ·  Molecular phylogenetic
investigation has shown the Zygomycota to be a polyphyletic phylum with evidence of
paraphyly within this taxonomic group

6c Members of the Glomeromycota are fungi forming arbuscular mycorrhizae with higher
plants Only one species has been observed forming zygospores; all other species solely
reproduce asexually The symbiotic association between the Glomeromycota and plants
is ancient, with evidence dating to  million years ago


Diagram of an apothecium (the typical cup-like reproductive structure of AscomycetesÚ showing

sterile tissues as well as developing and mature asci
 6c The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest
taxonomic group within the Eumycota These fungi form meiotic spores called
ascospores, which are enclosed in a special sac-like structure called an ascus This
division includes morels, a few mushrooms and truffles, single-celled yeasts (e g , of the
genera 

, -  , ]
, and 


Ú, and many filamentous
fungi living as saprotrophs, parasites, and mutualistic symbionts Prominent and
important genera of filamentous ascomycetes include   , ]
 , © 
 ,
and 
 Many ascomycetes species have only been observed undergoing asexual
reproduction (called anamorphic speciesÚ, but molecular data has often been able to
identify their closest teleomorphs in the Ascomycota Because the products of meiosis are
retained within the sac-like ascus, several ascomyctes have been used for elucidating
principles of genetics and heredity (e g % 


Ú
6c Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes,
produce meiospores called basidiospores on club-like stalks called basidia Most common
mushrooms belong to this group, as well as rust (fungusÚ and smut fungi, which are major
pathogens of grains Other important Basidiomyces include the maize pathogen,(


, human commensal species of the genus 0


, and the opportunistic human
pathogen,   
S 





 


  S
 
 

 

Because of some similarities in morphology and lifestyle, the slime molds (myxomycetesÚ and
water molds (oomycetesÚ were formerly classified in the kingdom Fungi Unlike true fungi,
however, the cell walls of these organisms contain cellulose and lack chitin Slime molds are
unikonts like fungi, but are grouped in the Amoebozoa Water molds are diploid bikonts,
grouped in the Chromalveolate kingdom Neither water molds nor slime molds are closely
related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi
Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology
textbooks and primary research literature

It has been suggested that the nucleariids, currently grouped in the Choanozoa, may be a sister
group to the oomycete clade, and as such could be included in an expanded fungal kingdom
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