we’ll see later in the chapter, cell walls made of chitin, and they also share certain key

one such chytrid parasite enzymes and metabolic pathways with other fungal groups.
has likely contributed to the Some chytrids form colonies with hyphae, while others exist
Flagellum
global decline of amphibian as single spherical cells.

4 μm
populations. Still other chy-
trids are important mutual-
ists. For example, anaerobic ▲ Figure 31.11  Flagellated
Zygomycetes
chytrid zoospore (TEM).
chytrids that live in the di- Chytrids There are approximately 1,000
gestive tracts of sheep and cattle help to break down plant Zygomycetes known species of zygomycetes,
Glomeromycetes
matter, thereby contributing significantly to the animal’s fungi in the phylum Zygomycota.
Ascomycetes
growth. Basidiomycetes This diverse phylum includes species
As discussed earlier, molecular evidence indicates that of fast-growing molds responsible
some chytrid lineages diverged early in fungal evolution. for causing foods such as bread, peaches, strawberries, and
The fact that chytrids are unique among fungi in having sweet potatoes to rot during storage. Other zygomycetes live
flagellated spores, called zoospores (Figure 31.11), agrees as parasites or as commensal (neutral) symbionts of animals.
with this hypothesis. Like other fungi, chytrids (other than The life cycle of Rhizopus stolonifer (black bread mold)
those in the recently discovered cryptomycota clade) have is fairly typical of zygomycete species (Figure 31.12). Its

1 Mycelia have various 2 Neighboring mycelia of different Key

mating types (here designated (–), mating types form hyphal extensions
with red nuclei, and (+), with (gametangia), each of which encloses Haploid (n)
blue nuclei). several haploid nuclei.
Heterokaryotic (n + n)
Diploid (2n)

3 A zygosporangium
PLASMOGAMY forms, containing
multiple haploid nuclei
Mating Gametangia with from the two parents.
type (–) haploid nuclei
Mating
type (+)

Rhizopus 100 μm
growing Young
on bread zygosporangium
(heterokaryotic)
8 The spores
germinate and
SEXUAL
grow into new
REPRODUCTION
9 Mycelia can also reproduce mycelia.
asexually by forming sporangia
that produce genetically
identical haploid spores. Dispersal and Zygosporangium
germination
KARYOGAMY
Sporangia
7 The sporangium 4 The zygosporangium
disperses genetically develops a rough,
diverse haploid spores. thick-walled coating
Diploid that can resist harsh
nuclei conditions for months.
Sporangium
ASEXUAL
REPRODUCTION MEIOSIS

Dispersal and 5 When conditions are

germination favorable, karyogamy
occurs, then meiosis.

50 μm Mycelium
6 The zygosporangium
▲ Figure 31.12  The life cycle of the zygomycete germinates into a
Rhizopus stolonifer (black bread mold). sporangium on a short stalk.

656    U n i t F i v e    The Evolutionary History of Biological Diversity

 Ascomycetes vary in size and complexity from unicellular
yeasts to elaborate cup fungi and morels (see Figure 31.15).
They include some of the most devastating plant pathogens,
which we will discuss later. However, many ascomycetes are
important decomposers, particularly of plant material. More
than 25% of all ascomycete species live with green algae or
cyanobacteria in beneficial symbiotic associations called
lichens. Some ascomycetes form mycorrhizae with plants.
Many others live between mesophyll cells in leaves; some of
these species release toxic compounds that help protect the
plant from insects.
Although the life cycles of various ascomycete groups
differ in the details of their reproductive structures and
processes, we’ll illustrate some common elements using the
bread mold Neurospora crassa (Figure 31.16). Ascomycetes
reproduce asexually by producing enormous numbers of
asexual spores called conidia (singular, conidium). Unlike
the asexual spores of most zygomycetes, conidia are not
formed inside sporangia. Rather, they are produced exter-
nally at the tips of specialized hyphae called conidiophores,
often in clusters or long chains, from which they may be
dispersed by the wind.
Conidia may also be involved in sexual reproduction,
fusing with hyphae from a mycelium of a different mat-
ing type, as occurs in Neurospora. Fusion of two different
mating types is followed by plasmogamy, resulting in the

1 Ascomycete mycelia 2 Neurospora can also reproduce Key

can reproduce asexually Conidia; sexually by producing specialized
by producing pigmented mating type (–) hyphae. Conidia of the opposite Haploid (n)
haploid spores (conidia). mating type fuse to these hyphae.
Dikaryotic (n + n)
Diploid (2n)

Dispersal
Germination Mating 3 The dikaryotic hyphae
type (+) that result from
ASEXUAL
plasmogamy produce
REPRODUCTION Hypha PLASMOGAMY many dikaryotic asci, two
of which are shown here.

Ascus
(dikaryotic)
Conidiophore
Mycelia
Dikaryotic
hyphae
Mycelium

Germination SEXUAL
REPRODUCTION KARYOGAMY
Dispersal
Diploid nucleus 4 Karyogamy
(zygote) occurs within each
Asci ascus, producing a
Eight
Ascocarp ascospores diploid nucleus.

7 The ascospores
are discharged forcibly
from the asci through an Four
opening in the ascocarp. haploid
Germinating ascospores nuclei
give rise to new mycelia. MEIOSIS

6 Each haploid nucleus divides once by

5 Each diploid nucleus
mitosis, yielding eight nuclei. Cell walls
and plasma membranes develop around divides by meiosis, yielding
the nuclei, forming ascospores (LM). four haploid nuclei.

▲ Figure 31.16  The life cycle of Neurospora crassa, an ascomycete. Neurospora is a bread
mold and research organism that also grows in the wild on burned vegetation.

658    U n i t F i v e    The Evolutionary History of Biological Diversity

 The life cycle of a basidiomycete usually includes a
long-lived dikaryotic mycelium (Figure 31.18). As in
ascomy­cetes, this extended dikaryotic stage provides many
opportunities for genetic recombination events, in effect
multiplying the result of a single mating. Periodically, in
response to environmental stimuli, the mycelium repro-
duces sexually by producing elaborate fruiting bodies called
basidiocarps. The common white mushrooms in the super-
market are familiar examples of a basidiocarp.
By concentrating growth in the hyphae of mushrooms, a
basidiomycete mycelium can erect its fruiting structures in
just a few hours; a mushroom pops up as it absorbs water
and as cytoplasm streams in from the dikaryotic mycelium.
By this process, a ring of mushrooms, popularly called a
“fairy ring,” may appear literally overnight (Figure 31.19).
The mycelium below the fairy ring expands outward at a
rate of about 30 cm per year, decomposing organic matter in
the soil as it grows. Some giant fairy rings are produced by
mycelia that are centuries old.
After a mushroom forms, its cap supports and protects
a large surface area of dikaryotic basidia on gills. During
karyogamy, the two nuclei in each basidium fuse, producing
a diploid nucleus (see Figure 31.18). This nucleus then un-
dergoes meiosis, yielding four haploid nuclei, each of which
ultimately develops into a basidiospore. Large numbers of
basidiospores are produced: The gills of a common white
mushroom have a surface area of about 200 cm2 and may
drop a billion basidiospores, which blow away.

▼ Figure 31.18  The life 2 A dikaryotic mycelium forms,

cycle of a mushroom- 1 Two haploid mycelia growing faster than, and ultimately
forming basidiomycete. of different mating types crowding out, the haploid parental mycelia.
undergo plasmogamy.

Dikaryotic
PLASMOGAMY mycelium 3 Environmental cues
such as rain or change in
temperature induce the
dikaryotic mycelium to
Mating form compact masses
8 In a suitable type (–) that develop into
environment, the basidiocarps (mushrooms,
basidiospores in this case).
germinate and Mating
grow into type (+)
short-lived
Haploid
haploid mycelia.
mycelia Gills lined
SEXUAL with basidia
REPRODUCTION Basidiocarp
(n + n)

7 When mature,
the basidiospores
Dispersal
are ejected and
and
then dispersed
germination
by the wind.

Basidiospores
(n)

Basidium with Basidia

four basidiospores (n + n)
Basidium

Basidium containing 4 The basidiocarp

four haploid nuclei gills are lined with
terminal dikaryotic
KARYOGAMY cells called basidia.

MEIOSIS

Key
6 Each diploid nucleus
yields four haploid 5 Karyogamy in each Haploid (n)
nuclei, each of which Diploid basidium produces a
1 μm Basidiospore develops into a nuclei diploid nucleus, which Dikaryotic (n + n)
basidiospore (SEM). then undergoes meiosis. Diploid (2n)

660    U n i t F i v e    The Evolutionary History of Biological Diversity