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19/8/13

Phylum Choanoagellata
The Choanoagellata is a small phylum,
containing only about 200 species of unicellular,
solitary, or colony-forming agellates without
chloroplasts.
The cells of some of the colonial species are
united by cytoplasmic bridges, which have not
been studied in detail, but which appear to be
the remains of incomplete cell divisions so that
the colonies are actually plasmodia; cell juncIons
have not been reported.

Phylum Choanoagellata

The colonies are plate-like


or spherical; the more well-
known spherical types, such
as Sphaeroeca (see the
chapter vigneLe), have the
collar complexes on the
outer side of the sphere,
but Diaphanoeca has the
collar complexes facing an
internal cavity, so that the
colonies resemble free-
swimming collar chambers
of a sponge.

Prelude:
Phylum Choanoflagellata
The Choanoflagellata is a small phylum, containing
only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelatinous envelope, but a large group of marine and
brackish-water species have an elaborate lorica consisting of siliceous costae united into an elegant bellshaped meshwork. The costae are secreted in small
vacuoles and transported to the surface of the cell,
where they become arranged in species-specific patterns (Leadbeater 1994).

Phylum Choanoagellata
Choanoagellates occur in
most aquaIc habitats and
are either pelagic or sessile.
The apparently most
primiIve species are naked
or sheathed in a gelaInous
envelope, but a large group
of marine and brackish-
water species have an
elaborate lorica consisIng
of siliceous costae united
into an elegant bellshaped
meshwork.

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

Prelude:
Phylum Choanoflagellata

Chapter vignette: Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)

The Choanoflagellata is a small phylum, containing


only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelati-

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and

19/8/13

Phylum Choanoagellata

The costae are secreted


in small vacuoles and
transported to the
surface of the cell,
where they become
arranged in species-
specic paLerns
(Leadbeater 1994).

Prelude:
Phylum Choanoflagellata
The Choanoflagellata is a small phylum, containing
only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelatinous envelope, but a large group of marine and
brackish-water species have an elaborate lorica consisting of siliceous costae united into an elegant bellshaped meshwork. The costae are secreted in small
vacuoles and transported to the surface of the cell,
where they become arranged in species-specific patterns (Leadbeater 1994).

Phylum Choanoagellata
The ovoid cell body has
a circle of 1550
microvilli or tentacles,
forming a funnel
surrounding a long
undulaIng cilium.

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

Prelude:
Phylum Choanoflagellata

Chapter vignette: Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)

The Choanoflagellata is a small phylum, containing


only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelatinous envelope, but a large group of marine and
brackish-water species have an elaborate lorica consisting of siliceous costae united into an elegant bellshaped meshwork. The costae are secreted in small
vacuoles and transported to the surface of the cell,
where they become arranged in species-specific patterns (Leadbeater 1994).

Phylum Choanoagellata
The microvilli are
retracIle and contain
acIn, and the whole
structure is held in
shape by a mucous or
brillar meshwork
(Karpov and Leadbeater
1998 ; King 2005).

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

Prelude:
Phylum Choanoflagellata

Chapter vignette: Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)

The Choanoflagellata is a small phylum, containing


only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelati-

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and

19/8/13

Phylum Choanoagellata

The single cilium (usually called a


agellum) is usually much longer
than the collar and has a dainty
extracellular brillar vane
(Leadbeater 2006).
The cilium forms sinusoidal waves
travelling towards the Ip of the
cilium, pumping water between
the microvilli; bacteria and other
food items are retained on the
outside of the collar and become
engulfed by pseudopodia formed
from the base of the collar
(Leadbeater 1977 , 1983a).
In pelagic species, the ciliary
acIvity is also locomotory.

Prelude:
Phylum Choanoflagellata
The Choanoflagellata is a small phylum, containing
only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelatinous envelope, but a large group of marine and
brackish-water species have an elaborate lorica consisting of siliceous costae united into an elegant bellshaped meshwork. The costae are secreted in small
vacuoles and transported to the surface of the cell,
where they become arranged in species-specific patterns (Leadbeater 1994).

Phylum Choanoagellata
The basal body of the
ciliary axoneme is
surrounded by an
intricate system of
radiaIng microtubules,
and there is an
accessory centriole that
is oriented at right
angles to the basal body
in some species (Karpov
and Leadbeater 1997).

Prelude:
Phylum Choanoflagellata

Chapter vignette: Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)

The Choanoflagellata is a small phylum, containing


only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelatinous envelope, but a large group of marine and
brackish-water species have an elaborate lorica consisting of siliceous costae united into an elegant bellshaped meshwork. The costae are secreted in small
vacuoles and transported to the surface of the cell,
where they become arranged in species-specific patterns (Leadbeater 1994).

Phylum Choanoagellata
ReproducIon is by
binary ssion.
Sexual reproducIon has
not been observed, but
the presence of
conserved meioIc
genes indicates that the
process has just gone
unnoIced (Carr et al.
2010).

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

Prelude:
Phylum Choanoflagellata

Chapter vignette: Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)

The Choanoflagellata is a small phylum, containing


only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelati-

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and

19/8/13

Phylum Choanoagellata

Also the presence of a


complicated life cycle,
with dierent cell and
colony types in one
species, (Leadbeater
1983b) indicates the
presence of sexual
reproducIon. The
ploidy level is unknown.

Prelude:
Phylum Choanoflagellata

The ovoid cell body has a circle of 1550 microThe Choanoflagellata is a small phylum, containing
villi or tentacles, forming a funnel surrounding a long
only about 200 species of unicellular, solitary, or
undulating cilium. The microvilli are retractile and
colony-forming flagellates without chloroplasts. The
contain actin, and the whole structure is held in shape
cells of some of the colonial species are united by cytoby a mucous or fibrillar meshwork (Karpov and
plasmic bridges, which have not been studied in detail,
Leadbeater 1998; King 2005). The single cilium (usubut which appear to be the remains of incomplete cell
ally called a flagellum) is usually much longer than
divisions so that the colonies are actually plasmodia;
the collar and has a dainty extracellular fibrillar vane
cell junctions have not been reported. The colonies
(Leadbeater 2006). The cilium forms sinusoidal waves
areEvolution
plate-like or spherical; the more well-known
4
1 Animal
travelling towards the tip of the cilium, pumping
spherical types, such as
Sphaeroeca (see the chapter
system after a period with the popular classification of
(Hejnol et al. 2009), and analyses based on whole
Bilateria = Acoela + Pseudocoela + Coelomata, which
genomes, (Srivastava et al. 2008, 2010), now seem to
water between the microvilli; bacteria and other food
vignette),
have
the
collar
complexes
on
the
outer
side
is a misunderstanding of the presentation of Hyman
bring much more stability to the more basal parts of
(1940) (see Nielsen 2010).
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
items are retained on the outside of the collar and
of the sphere, but Diaphanoeca
has the collar complexes
It should also be remembered that the molecular
and microRNAs appear to be of special value in the
phylogenies are naked trees without morphological
studies of evolution of the major metazoan groups
become engulfed by pseudopodia formed from the
facing
an
internal
cavity,
so
that
the
colonies
resemble
characters, and, as stated by Raff et al. (1989, p. 258),
(de Rosa et al. 1999; Peterson et al. 2009).
The use of rRNA sequences to infer distant phylogeIt should be emphasized that the classification of
base of the collar (Leadbeater 1977, 1983a). In pelagic
free-swimming
collar
chambers
of
a
sponge.
The
netic relationships will not displace morphology and
Bilaterians in Protostomia and Deuterostomia dates
embryology
from
the
study
of
the
evolutionary
history
back to Grobben
(
1908
),
so
it
is
not
the
new
phylogspecies, the ciliary activity is also locomotory. The
monophyly of the group
seems unquestioned (Carr
of animal life: after all, it is the history of morphological
eny as proposed, especially in a number of papers on
molecular phylogenetics,
but a return
basal body of the
ciliary axoneme is surrounded by an
et al. 2008
). to the classical change that we wish to explain [my italics].
Prelude:
intricate
systemagellat
of radiating
microtubules, and there
Choanoflagellates occur in most aquatic habitats
Phylum
Choanofl
a
A - Animalia
Vertebrata
Urochordata
is
an
accessory
centriole
that is oriented at right angles
B - Euradiculata
and
are either pelagic or sessile.
The
apparently
most
Z
Cephalochordata
C - Proepitheliozoa
The ovoid cell body has a circle of 1550 microThe Choanofl
agellata is a small phylum, containing
V
D - Eumetazoa
villi
a funnelbody
surroundingin
a long some species (Karpov and
about
200 species of
solitary, or
toor tentacles,
the forming
basal
primitive
species are naked
or Pterobranchia
sheathed
inunicellular,
a gelatiEnteropneusta
X Y only
E - Neuralia
Echinodermata
undulating cilium. The microvilli are retractile and
colony-forming
flagellates without chloroplasts. The
F - Triploblastica
contain
actin, and the whole1997
structure).
is held in shape
ofChaetognatha
some of theof
colonial
species are united
by cytoLeadbeater
nous
envelope, but a largecellsgroup
marine
and
G - Bilateria
Loricifera
by a mucous or fibrillar meshwork (Karpov and
bridges, which have not been studied in detail,
H- Acoelomorpha
Kinorhyncha
Uplasmic
I - Eubilateria
I
1998; King 2005). The single
but which
appear to be the remains
of incomplete
Priapula
S an
Reproduction
iscilium
by (usubinary fission. Sexual reproducbrackish-water
species
have
elaborate
lorica
con-cell Leadbeater
K - Protostomia
Nematomorpha
ally called a flagellum) is usually much longer than
so that the colonies are actually plasmodia;
T divisions
Nematoda
L - Spiralia
Q
thetion
collar and
has a dainty
extracellular
fi
brillar vane
cell junctions
not been
reported. The
colonies
Tardigrada
has
not
been
observed,
but the presence of consisting
of siliceous costae united
intohave
an
elegant
bellM - Schizocoelia
Onychophora
R
(
Leadbeater
2006
).
The
cilium
forms
sinusoidal
waves
are
plate-like
or
spherical;
the
more
well-known
N - Gnathifera
Arthropoda
travelling towards the tip of the cilium, pumping
spherical types, such as Sphaeroeca (see the chapter
O - Polyzoa
served meiotic genes indicates that the process has just
Brachiopoda
shaped
meshwork. The
costae
are
secreted in small
P vignette),
K
P - Brachiozoa
water between the microvilli; bacteria and other food
have the collar complexes on the outer side
Phoronida
Q - Ecdysozoa
items are retained on the outside of the collar and
of theBryozoa
sphere, but Diaphanoeca has the collar complexes
gone
unnoticed
(
Carr
et
al.
2010
). Also the presence of
vacuoles
and transportedOto facing
the
surface
of
the
cell,
R - Panarthropoda
Cycliophora
become engulfed by pseudopodia formed from the
an internal cavity, so that the colonies resemble
Entoprocta
S - Cycloneuralia
G
base of the collar (Leadbeater 1977, 1983a). In pelagic
free-swimming
collar chambers of a sponge. The
T - Nematoidea
a complicated life cycle, with different cell and colony
where
they become arranged
inRotifera
species-specifi
c patMicrognathozoa
species, the ciliary activity is also locomotory. The
of the group seems unquestioned (Carr
N monophyly
U - Scalidophora
Gnathostomulida
L
V - Deuterostomia
basal body of the ciliary axoneme is surrounded by an
et al. 2008
).
Gastrotricha
types
in
one
species,
(
Leadbeater
1983b) indicates the
terns
(Leadbeater 1994).
X - Ambulacraria
intricate system of radiating microtubules, and there
Choanofl
agellates occur in most aquatic habitats
Platyhelminthes

Phylum Choanoagellata
The sister-group
relaIonship of
choanoagellates and
metazoans is supported
by numerous
morphological and
molecular analyses,
including analyses of
whole genomes (King et
al. 2008).

Y - Hemichordata
Z - Chordata

F
E

M
H

Nemertini
and are
either pelagic or sessile. The apparently most
Mollusca
primitive
species are naked or sheathed in a gelatiSipuncula
nous Annelida
envelope, but a large group of marine and
Xenoturbellida
brackish-water
species have an elaborate lorica conNemertodermatida
sistingAcoela
of siliceous costae united into an elegant bellshaped
meshwork. The costae are secreted in small
Ctenophora
Cnidaria
vacuoles
and transported to the surface of the cell,
Placozoa
whereHomoscleromorpha
they become arranged in species-specific patterns Calcarea
(Leadbeater 1994).

is an accessory centriole that is oriented at right angles


to the basal body in some species (Karpov and
Leadbeater 1997).
Reproduction is by binary fission. Sexual reproduction has not been observed, but the presence of conserved meiotic genes indicates that the process has just
gone unnoticed (Carr et al. 2010). Also the presence of
a complicated life cycle, with different cell and colony
types in one species, (Leadbeater 1983b) indicates the

Chapter vignette:
Sphaeroeca volvox. (Redrawn from Leadbeater 1983b.)
D
A

Silicea
Choanoflagellata

Figure 1.2. The animal phylogeny proposed in this book; the method is described
in thevignette:
text. Named
supraphyletic
clades are
Chapter
Sphaeroeca
volvox. (Redrawn
from Leadbeater 1983b.)
indicated.

Phylum Choanoagellata
Many parts of the
molecular machinery
involved in
mulIcellularity are
present, but with
unknown funcIons and
have supposedly
become co-opted into
their cellcell
interacIons in the
metazoans (King 2004).

Prelude:
Phylum Choanoflagellata
The Choanoflagellata is a small phylum, containing
only about 200 species of unicellular, solitary, or
colony-forming flagellates without chloroplasts. The
cells of some of the colonial species are united by cytoplasmic bridges, which have not been studied in detail,
but which appear to be the remains of incomplete cell
divisions so that the colonies are actually plasmodia;
cell junctions have not been reported. The colonies
are plate-like or spherical; the more well-known
spherical types, such as Sphaeroeca (see the chapter
vignette), have the collar complexes on the outer side
of the sphere, but Diaphanoeca has the collar complexes
facing an internal cavity, so that the colonies resemble
free-swimming collar chambers of a sponge. The
monophyly of the group seems unquestioned (Carr
et al. 2008).
Choanoflagellates occur in most aquatic habitats
and are either pelagic or sessile. The apparently most
primitive species are naked or sheathed in a gelati-

The ovoid cell body has a circle of 1550 microvilli or tentacles, forming a funnel surrounding a long
undulating cilium. The microvilli are retractile and
contain actin, and the whole structure is held in shape
by a mucous or fibrillar meshwork (Karpov and
Leadbeater 1998; King 2005). The single cilium (usually called a flagellum) is usually much longer than
the collar and has a dainty extracellular fibrillar vane
(Leadbeater 2006). The cilium forms sinusoidal waves
travelling towards the tip of the cilium, pumping
water between the microvilli; bacteria and other food
items are retained on the outside of the collar and
become engulfed by pseudopodia formed from the
base of the collar (Leadbeater 1977, 1983a). In pelagic
species, the ciliary activity is also locomotory. The
basal body of the ciliary axoneme is surrounded by an
intricate system of radiating microtubules, and there
is an accessory centriole that is oriented at right angles
to the basal body in some species (Karpov and

19/8/13

Phylum Choanoagellata

The Six Steps!


FIRST MAJOR STEP: THE EVOLUTION OF
MULTICELLULARITY (METAZOA)
MOLECULAR AND COMBINED STUDIES OF
EARLY METAZOAN RADIATION (TABLE 1)
EVOLUTION OF THE EARLIEST METAZOANS,
THE SPONGES

SECOND MAJOR STEP: THE ORIGIN OF SEALED


EPITHELIA AND EXTRACELLULAR DIGESTION
(EUMETAZOA)
PLACOZOA: Trichoplax adherens

19/8/13

THIRD MAJOR STEP: THE ORIGIN OF A


NERVOUS SYSTEM (NEURALIA)
CNIDARIA

FOURTH MAJOR STEP: THE ORIGIN OF


MESODERM (TRIPLOBLASTICA)
CTENOPHORA

FIFTH MAJOR STEP: THE ORIGIN OF


BILATERALITY (BILATERIA)
SIXTH MAJOR STEP: THE ESTABLISHMENT OF
A TUBULAR GUT (EUBILATERIA)

19/8/13

Phylum Choanoagellata

4 Animal Evolution

16

2 m

5 m

5m

Figure 4.1. SEM of a choanoflagellate and collar chambers of two types of sponges. (A) A solitary stage of the choanoflagellate
Salpingoeca sp. (courtesy of Drs M.J. Dayel and N. King, University of California, Berkeley, CA, USA.) (B) The demosponge
Callyspongia diffusa (courtesy of Dr I.S. Johnston, Bethel College, MI, USA; see Johnston and Hildemann 1982). (C) The calcarean Sycon sp. (Friday Harbor Laboratories, WA, USA, July 1988.)

topologies presented by these papers indicate that


these genes are not sufficiently informative for resolving the deep-branching pattern of the metazoans. A
further reason for this is that the taxon sampling has
only recently begun to include sponges other than
demosponges.
The ancestral metazoan was probably a choanoblastaea (Fig. 4.3; see also Chapter 2 and Nielsen
2008), which had developed so close cell contacts that
nutrients could be exchanged between cells; this made
specialization of cells possible, because not all cells
had to feed. It probably had a life cycle with sexual
reproduction and haploid eggs and sperm, characteristic of the living metazoans. Sexual reproduction originated at an early step in the evolution of eukaryotes
(Dacks and Roger 1999), but it could of course have
been lost in the choanoflagellates.
The new understanding of the early metazoan
radiation with a paraphyletic Porifera implies that the
life cycle found in the sponge grade, with a ciliated

free-swimming larva and a sessile adult form with a


water-canal system and choanocyte chambers, was
ancestral. The evolution of this type of organization is
visualized in Fig. 4.3.
A choanoblastaea lineage developed a number of
internalized cells (the advanced choanoblastaea), and
established a polarity with special cells without collars
at the anterior pole. These cells were used in settling,
and the choanocytes of the benthic stage became
arranged in a longitudinal groove, which established a
common water current along the groove, enhancing
the current and preventing recirculation of already
processed water. This ancestral sponge retained the
pelagic stage as a dispersal larval stage, and a pelagobenthic life cycle had become established. The groove
with the vulnerable choanocytes became protected by
overarching from the lateral cells so that a tubular
choanocyte chamber was formed, further enhancing
the water currents (the primitive sponge). This is an
organization resembling the living calcareans of the

Origen de Metazoa

2 teoras origen metazoarios

Teora colonial-
Haeckel
Teora celularizacin
(sincicial)-Hadzi,
Steinbck

19/8/13

2 teoras origen metazoarios


Teora colonial-Haeckel

Teora celularizacin (sincicial)-


Hadzi, Steinbck

The colonial theory proposes that the


rst mulIcellular organism evolved
from a colony of cells, derived from a
zygote, which developed cell contacts
enabling exchange of nutrients
between the cells and, subsequently,
specializaIon of cells.
Synamoebium - amoeboid cells that
later became ciliated (Haeckel 1868)

2 teoras origen metazoarios


Teora colonial-Haeckel

Ciliated hollow blastaea (Haeckel 1875).


Colonial agellates Volvox and Synura
as living examples of organisms showing
the same type of organizaIon (Haeckel
1889)
Metschniko (1886) seems to have
been the rst to discuss a
choanoagellate origin of the
metazoans (in the light of the newly
described Proterospongia ). The
quesIon moved out of focus for more
than half a century, but Remane (1963)
argued explicitly for a spherical
choano agellate colony as an ancestor
of the monophyleIc Metazoa.

Volvox

2 teoras origen metazoarios


Teora colonial-Haeckel

A sister-group relaIonship between


choanoagellates is supported both
by the detailed similarity between
choanoagellates and the
choanocytes of the sponges ( Fig.
4.1 ), and with several newer
molecular phylogenies (Chapter 3).
These results have been combined
and presented in a previous paper
(Nielsen 2008), and will be the
backbone of the phylogeny explained
below and in the following chapters.

Teora celularizacin (sincicial)-


Hadzi, Steinbck

The alternaIve
cellularizaIon theories,
which derive a
turbellariform-metazoan
ancestor through
compartmentalizaIon of a
ciliate, or a ciliate-like
organism (see e.g. Hadzi
1953 ; Steinbck 1963 ), is
now only of historical
interest.

19/8/13

1 Animal Evolution
system after a period with the popular classification of
Bilateria = Acoela + Pseudocoela + Coelomata, which
is a misunderstanding of the presentation of Hyman
(1940) (see Nielsen 2010).
It should also be remembered that the molecular
phylogenies are naked trees without morphological
characters, and, as stated by Raff et al. (1989, p. 258),
The use of rRNA sequences to infer distant phylogenetic relationships will not displace morphology and
embryology from the study of the evolutionary history
of animal life: after all, it is the history of morphological
change that we wish to explain [my italics].

(Hejnol et al. 2009), and analyses based on whole


genomes, (Srivastava et al. 2008, 2010), now seem to
bring much more stability to the more basal parts of
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
and microRNAs appear to be of special value in the
studies of evolution of the major metazoan groups
(de Rosa et al. 1999; Peterson et al. 2009).
It should be emphasized that the classification of
Bilaterians in Protostomia and Deuterostomia dates
back to Grobben (1908), so it is not the new phylogeny as proposed, especially in a number of papers on
molecular phylogenetics, but a return to the classical

2 teoras origen metazoarios

Teora celularizacin (sincicial)-


Hadzi, Steinbck

Teora colonial-Haeckel

Monophyly of porifera is no more


A - Animalia
B - Euradiculata
C - Proepitheliozoa
D - Eumetazoa
E - Neuralia
F - Triploblastica
G - Bilateria
H- Acoelomorpha
I - Eubilateria
K - Protostomia
L - Spiralia
M - Schizocoelia
N - Gnathifera
O - Polyzoa
P - Brachiozoa
Q - Ecdysozoa
R - Panarthropoda
S - Cycloneuralia
T - Nematoidea
U - Scalidophora
V - Deuterostomia
X - Ambulacraria
Y - Hemichordata
Z - Chordata

Vertebrata
Urochordata
Cephalochordata
Pterobranchia
Enteropneusta
Echinodermata
Chaetognatha
Loricifera
Kinorhyncha
Priapula
Nematomorpha
Nematoda
Tardigrada
Onychophora
Arthropoda
Brachiopoda
Phoronida
Bryozoa
Cycliophora
Entoprocta
Rotifera
Micrognathozoa
Gnathostomulida
Gastrotricha
Platyhelminthes
Nemertini
Mollusca
Sipuncula
Annelida
Xenoturbellida
Nemertodermatida
Acoela
Ctenophora
Cnidaria
Placozoa
Homoscleromorpha
Calcarea
Silicea
Choanoflagellata

Z
V

X Y

S
Q

U
T

R
K

P
O

G
L
F

D
A

The alternaIve
cellularizaIon theories,
which derive a
turbellariform-metazoan
ancestor through
compartmentalizaIon of a
ciliate, or a ciliate-like
organism (see e.g. Hadzi
1953 ; Steinbck 1963 ), is
now only of historical
interest.

Figure 1.2. The animal phylogeny proposed in this book; the method is described in the text. Named supraphyletic clades are
indicated.

2 teoras origen metazoarios


Recent molecular studies (for example,
Sperling et al. 2007 , 2009 ; Peterson et al .
2008 ), and a few morphological studies
(Cavalier-Smith et al. 1996 ; Borchiellini et al.
2001 ) have cast doubt about the monophyly
of the Porifera, and indicated a phylogeny like
that shown in Fig. 4.2 (Nielsen 2008).

1 Animal Evolution

(Hejnol et al. 2009), and analyses based on whole


genomes, (Srivastava et al. 2008, 2010), now seem to
bring much more stability to the more basal parts of
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
and microRNAs appear to be of special value in the
studies of evolution of the major metazoan groups
(de Rosa et al. 1999; Peterson et al. 2009).
It should be emphasized that the classification of
Bilaterians in Protostomia and Deuterostomia dates
back to Grobben (1908), so it is not the new phylogeny as proposed, especially in a number of papers on
molecular phylogenetics, but a return to the classical

system after a period with the popular classification of


Bilateria = Acoela + Pseudocoela + Coelomata, which
is a misunderstanding of the presentation of Hyman
(1940) (see Nielsen 2010).
It should also be remembered that the molecular
phylogenies are naked trees without morphological
characters, and, as stated by Raff et al. (1989, p. 258),
The use of rRNA sequences to infer distant phylogenetic relationships will not displace morphology and
embryology from the study of the evolutionary history
of animal life: after all, it is the history of morphological
change that we wish to explain [my italics].

2 teoras origen metazoarios

A - Animalia
B - Euradiculata
C - Proepitheliozoa
D - Eumetazoa
E - Neuralia
F - Triploblastica
G - Bilateria
H- Acoelomorpha
I - Eubilateria
K - Protostomia
L - Spiralia
M - Schizocoelia
N - Gnathifera
O - Polyzoa
P - Brachiozoa
Q - Ecdysozoa
R - Panarthropoda
S - Cycloneuralia
T - Nematoidea
U - Scalidophora
V - Deuterostomia
X - Ambulacraria
Y - Hemichordata
Z - Chordata

Z
V

S
Q

U
T

R
K

P
O

G
L
F
E

D
C

X Y

M
H

Vertebrata
Urochordata
Cephalochordata
Pterobranchia
Enteropneusta
Echinodermata
Chaetognatha
Loricifera
Kinorhyncha
Priapula
Nematomorpha
Nematoda
Tardigrada
Onychophora
Arthropoda
Brachiopoda
Phoronida
Bryozoa
Cycliophora
Entoprocta
Rotifera
Micrognathozoa
Gnathostomulida
Gastrotricha
Platyhelminthes
Nemertini
Mollusca
Sipuncula
Annelida
Xenoturbellida
Nemertodermatida
Acoela
Ctenophora
Cnidaria
Placozoa
Homoscleromorpha
Calcarea
Silicea
Choanoflagellata

Deuterostomia

Ecdysozoa

Lophotrochozoa Spiralia

Acoela
porifera

Figure 1.2. The animal phylogeny proposed in this book; the method is described in the text. Named supraphyletic clades are
indicated.

19/8/13

1 Animal Evolution

(Hejnol et al. 2009), and analyses based on whole


genomes, (Srivastava et al. 2008, 2010), now seem to
bring much more stability to the more basal parts of
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
and microRNAs appear to be of special value in the
studies of evolution of the major metazoan groups
(de Rosa et al. 1999; Peterson et al. 2009).
It should be emphasized that the classification of
Bilaterians in Protostomia and Deuterostomia dates
back to Grobben (1908), so it is not the new phylogeny as proposed, especially in a number of papers on
molecular phylogenetics, but a return to the classical

system after a period with the popular classification of


Bilateria = Acoela + Pseudocoela + Coelomata, which
is a misunderstanding of the presentation of Hyman
(1940) (see Nielsen 2010).
It should also be remembered that the molecular
phylogenies are naked trees without morphological
characters, and, as stated by Raff et al. (1989, p. 258),
The use of rRNA sequences to infer distant phylogenetic relationships will not displace morphology and
embryology from the study of the evolutionary history
of animal life: after all, it is the history of morphological
change that we wish to explain [my italics].

2 teoras origen metazoarios


This evoluIonary
scenario is based on a
combinaIon of
molecular data and
studies on embryology
and ultrastructure,
combined with
consideraIons of the
possible adapIve value
of each of the proposed
evoluIonary steps.

A - Animalia
B - Euradiculata
C - Proepitheliozoa
D - Eumetazoa
E - Neuralia
F - Triploblastica
G - Bilateria
H- Acoelomorpha
I - Eubilateria
K - Protostomia
L - Spiralia
M - Schizocoelia
N - Gnathifera
O - Polyzoa
P - Brachiozoa
Q - Ecdysozoa
R - Panarthropoda
S - Cycloneuralia
T - Nematoidea
U - Scalidophora
V - Deuterostomia
X - Ambulacraria
Y - Hemichordata
Z - Chordata

Z
V

X Y

S
Q

U
T

R
K

P
O

G
L
F

D
A

Vertebrata
Urochordata
Cephalochordata
Pterobranchia
Enteropneusta
Echinodermata
Chaetognatha
Loricifera
Kinorhyncha
Priapula
Nematomorpha
Nematoda
Tardigrada
Onychophora
Arthropoda
Brachiopoda
Phoronida
Bryozoa
Cycliophora
Entoprocta
Rotifera
Micrognathozoa
Gnathostomulida
Gastrotricha
Platyhelminthes
Nemertini
Mollusca
Sipuncula
Annelida
Xenoturbellida
Nemertodermatida
Acoela
Ctenophora
Cnidaria
Placozoa
Homoscleromorpha
Calcarea
Silicea
Choanoflagellata

Figure 1.2. The animal phylogeny proposed in this book; the method is described in the text. Named supraphyletic clades are
indicated.

1 Animal Evolution

(Hejnol et al. 2009), and analyses based on whole


genomes, (Srivastava et al. 2008, 2010), now seem to
bring much more stability to the more basal parts of
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
and microRNAs appear to be of special value in the
studies of evolution of the major metazoan groups
(de Rosa et al. 1999; Peterson et al. 2009).
It should be emphasized that the classification of
Bilaterians in Protostomia and Deuterostomia dates
back to Grobben (1908), so it is not the new phylogeny as proposed, especially in a number of papers on
molecular phylogenetics, but a return to the classical

system after a period with the popular classification of


Bilateria = Acoela + Pseudocoela + Coelomata, which
is a misunderstanding of the presentation of Hyman
(1940) (see Nielsen 2010).
It should also be remembered that the molecular
phylogenies are naked trees without morphological
characters, and, as stated by Raff et al. (1989, p. 258),
The use of rRNA sequences to infer distant phylogenetic relationships will not displace morphology and
embryology from the study of the evolutionary history
of animal life: after all, it is the history of morphological
change that we wish to explain [my italics].

2 teoras origen metazoarios


The scenario is in full accordance
with the phylogeny inferred by
Degnan et al. (2009), based on
transcripIon factors of the
choanoagellate Monosiga , the
siliceous sponge Amphimedon ,
Trichoplax , cnidarians
(Nematostella, Hydra), and
bilaterians, and with the whole-
genome studies of Srivastava et
al. (2008 , 2010),based on
Neurospora , Monosiga ,
Amphimedon , Trichoplax ,
Nematostella, Hydra , and a
number of bilaterians.

A - Animalia
B - Euradiculata
C - Proepitheliozoa
D - Eumetazoa
E - Neuralia
F - Triploblastica
G - Bilateria
H- Acoelomorpha
I - Eubilateria
K - Protostomia
L - Spiralia
M - Schizocoelia
N - Gnathifera
O - Polyzoa
P - Brachiozoa
Q - Ecdysozoa
R - Panarthropoda
S - Cycloneuralia
T - Nematoidea
U - Scalidophora
V - Deuterostomia
X - Ambulacraria
Y - Hemichordata
Z - Chordata

Z
V

X Y

S
Q

U
T

R
K

P
O

G
L
F

D
A

Vertebrata
Urochordata
Cephalochordata
Pterobranchia
Enteropneusta
Echinodermata
Chaetognatha
Loricifera
Kinorhyncha
Priapula
Nematomorpha
Nematoda
Tardigrada
Onychophora
Arthropoda
Brachiopoda
Phoronida
Bryozoa
Cycliophora
Entoprocta
Rotifera
Micrognathozoa
Gnathostomulida
Gastrotricha
Platyhelminthes
Nemertini
Mollusca
Sipuncula
Annelida
Xenoturbellida
Nemertodermatida
Acoela
Ctenophora
Cnidaria
Placozoa
Homoscleromorpha
Calcarea
Silicea
Choanoflagellata

Figure 1.2. The animal phylogeny proposed in this book; the method is described in the text. Named supraphyletic clades are
indicated.

1 Animal Evolution

(Hejnol et al. 2009), and analyses based on whole


genomes, (Srivastava et al. 2008, 2010), now seem to
bring much more stability to the more basal parts of
the phylogenetic tree. Both Hox genes (see Fig. 21.3)
and microRNAs appear to be of special value in the
studies of evolution of the major metazoan groups
(de Rosa et al. 1999; Peterson et al. 2009).
It should be emphasized that the classification of
Bilaterians in Protostomia and Deuterostomia dates
back to Grobben (1908), so it is not the new phylogeny as proposed, especially in a number of papers on
molecular phylogenetics, but a return to the classical

system after a period with the popular classification of


Bilateria = Acoela + Pseudocoela + Coelomata, which
is a misunderstanding of the presentation of Hyman
(1940) (see Nielsen 2010).
It should also be remembered that the molecular
phylogenies are naked trees without morphological
characters, and, as stated by Raff et al. (1989, p. 258),
The use of rRNA sequences to infer distant phylogenetic relationships will not displace morphology and
embryology from the study of the evolutionary history
of animal life: after all, it is the history of morphological
change that we wish to explain [my italics].

2 teoras origen metazoarios


However, it is not in agreement
with the numerous studies based
on ribosomal genes or various
collecIons of other smaller genes
but the very incongruent
topologies presented by these
papers indicate that these genes
are not suciently informaIve
for resolving the deep-branching
paLern of the metazoans. A
further reason for this is that the
taxon sampling has only recently
begun to include sponges other
than demosponges.

A - Animalia
B - Euradiculata
C - Proepitheliozoa
D - Eumetazoa
E - Neuralia
F - Triploblastica
G - Bilateria
H- Acoelomorpha
I - Eubilateria
K - Protostomia
L - Spiralia
M - Schizocoelia
N - Gnathifera
O - Polyzoa
P - Brachiozoa
Q - Ecdysozoa
R - Panarthropoda
S - Cycloneuralia
T - Nematoidea
U - Scalidophora
V - Deuterostomia
X - Ambulacraria
Y - Hemichordata
Z - Chordata

Z
V

S
Q

U
T

R
K

P
O

G
L
F
E

D
B

X Y

M
H

Vertebrata
Urochordata
Cephalochordata
Pterobranchia
Enteropneusta
Echinodermata
Chaetognatha
Loricifera
Kinorhyncha
Priapula
Nematomorpha
Nematoda
Tardigrada
Onychophora
Arthropoda
Brachiopoda
Phoronida
Bryozoa
Cycliophora
Entoprocta
Rotifera
Micrognathozoa
Gnathostomulida
Gastrotricha
Platyhelminthes
Nemertini
Mollusca
Sipuncula
Annelida
Xenoturbellida
Nemertodermatida
Acoela
Ctenophora
Cnidaria
Placozoa
Homoscleromorpha
Calcarea
Silicea
Choanoflagellata

Figure 1.2. The animal phylogeny proposed in this book; the method is described in the text. Named supraphyletic clades are
indicated.

10

19/8/13

2 teoras origen metazoarios


A

The ancestral metazoan was probably


a choanoblastaea (Fig. 4.3), which
had developed so close cell contacts
that nutrients could be exchanged
between cells; this made
specializaIon of cells possible,
because not all cells had to feed.
It probably had a life cycle with
sexual reproducBon and haploid
eggs and sperm, characterisIc of the
living metazoans.
Sexual reproducIon originated at an
early step in the evoluIon of
eukaryotes, but it could of course
have been lost in the
choanoagellates.

Choanoflagellate Choanoblastaea

Advanced
choanoblastaea

mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

2 teoras origen metazoarios


The new understanding of the early
metazoan radiaIon with a
paraphyleIc Porifera implies that the
life cycle found in the sponge grade,
with a ciliated free-swimming larva
and a sessile adult form with a water-
canal system and choanocyte
chambers, was ancestral. The
evoluIon of this type of organizaIon
visualized in Fig. 4.3. Advanced
Choanoflagellate is Choanoblastaea
choanoblastaea

mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

2 teoras origen mGastraea


etazoarios

Neurogastraea

A choanoblastaea lineage developed


a number of internalized cells (the
advanced choanoblastaea),
and
Homoscleromorph-like
Modern
established a pancestor
olarity with
withspecial
gastrula
homoscleromorph
cells without clarva
ollars and
at the
a
nterior
dissogony
pole.

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Choanoflagellate
Choanoblastaea
Plate 1. The evolutionary scenario (see text). The water currents are indicated
with blue arrows,
the basement Advanced
membrane is
choanoblastaea
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page
18.
mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

11

19/8/13

2 teoras origen metazoarios

A choanoblastaea lineage developed


A of internalized cells (the
a number
advanced choanoblastaea), and
established a polarity with special
cells without collars at the anterior
pole.
These cells were used in seLling, and
Choanoflagellate
the choanocytes
of the Choanoblastaea
benthic stage
became arranged in a longitudinal
groove, which
established
common
water a
current
mitosis
water current along the
groove,
basement
germ cells
enhancing the current membrane
and prevenIng
cadherins
junction
recirculaIon
of already occluding
processed
water.

Advanced
choanoblastaea
nerve
sensory cell
circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

2 teoras origen metazoarios

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

This ancestral sponge retained the


pelagic stage as a dispersal larval
stage, and a pelagobenthic life cycle
had become established.
The groove with the vulnerable
choanocytes became protected by
overarching from the lateral cells so
Choanoflagellate that
Choanoblastaea
Advanced
a tubular choanocyte
chamber
choanoblastaea
was formed, further enhancing
the
water currents (the primiIve
water
current
nerve
mitosis
sponge).

germ cells

basement
membrane

cadherins

occluding junction

sensory cell
circumblastoporal
nerve

Ancestral sponge

Primitive sponge
Ipo
Ascon

Gastraea
Neurogastraea
2 teoras origen metazoarios
A

This is an organizaIon resembling the

This grade is

This grade is

living calcareans
of the Ascon -type
represented
Homoscleromorph-like
represented
Modern
by Trichoplax
(see Chapter 7ancestor
), except with
that igastrula
t had no
by Cnidaria
homoscleromorph
skeleton. larva and dissogony
This form could grow to a larger size
Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
by organizing the layer of
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.
choanocytes into numerous separate
Choanoflagellate choanocyte
Choanoblastaea
chambers, Advanced
and the larger
size made it possible to choanoblastaea
invest more
yolk in the eggs, so that development
water
current
nerve
mitosis
could
become lecithotrophic.
germ cells

basement
membrane

cadherins

occluding junction

sensory cell
circumblastoporal
nerve

Ancestral sponge

Primitive sponge
Ascon

Gastraea
Neurogastraea

Modern

Homoscleromorph-like
ancestor with gastrula

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

12

19/8/13

2 teoras origen metazoarios


A

These non-feeding larvae abandoned


the external choanocytes with the
undulatory cilium, but developed a
new type of locomotory cilium, the
eecBve-stroke cilium.
This type of cilia has a long, more or
less organized root, and shows
coordinated movements between
neighbouring cells, creaIng
characterisIc metachronal waves*
(Fig. 2.1). This is the life cycle seen in
most living sponges.
* advance and recovery cycles

Choanoflagellate Choanoblastaea

Advanced
choanoblastaea

mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

Ascon

2 teoras origen metazoarios


A

Two lineages can be recognized from


the ancestral sponge: one lineage
leading to the Silicea, which
developed a siliceous skeleton
(Chapter 5); and another lineage
leading to the Euradiculata (Calcarea
+ Proepitheliozoa; Chapter 6 ) that
developed cross-striated ciliary roots
in the epithelium-like outer cell layer
of the larvae (Fig. 6.1). The
Calcareans developed calcareous
spicules ( Chapter 7 ).

Choanoflagellate Choanoblastaea

Advanced
choanoblastaea

mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

Ascon

2 teoras origen metazoarios


A

The Proepitheliozoa
(Homoscleromorpha + Eumetazoa)
developed collagen IV in the basal
membrane, and the rst stages of
occluding juncBons became
established.
One lineage led to the
Homoscleromorpha (Chapter 9),
which remained at this stage. The
other lineage led to the Eumetazoa,
which will be discussed in Chapter
10 .

Choanoflagellate Choanoblastaea

Advanced
choanoblastaea

mitosis

water current

nerve

germ cells

basement
membrane

sensory cell

cadherins

occluding junction

circumblastoporal
nerve

Ancestral sponge

Primitive sponge

Gastraea
Neurogastraea

Modern
homoscleromorph

Homoscleromorph-like
ancestor with gastrula
larva and dissogony

This grade is
represented
by Trichoplax

This grade is
represented
by Cnidaria

Plate 1. The evolutionary scenario (see text). The water currents are indicated with blue arrows, the basement membrane is
red, and the nervous cells green. The muscle cells of the gastraea have been omitted for clarity. See Figure 4.3, page 18.

Ascon

13

19/8/13

Descendemos de coanoagelados
coloniales

Denicin de Animal
NutriBonal Mode

Animals dier from both plants and fungi in


their mode of nutriIon.
Plants are autotrophic eukaryotes capable of
generaIng organic molecules through
photosynthesis.
Fungi are heterotrophs that grow on or near
their food and that feed by absorpBon (oyen
ayer they have released enzymes that digest
the food outside their bodies).
Unlike plants, animals cannot construct all of
their own organic molecules and so, in most
cases.
They ingest them-either by eaBng other
living organisms or by eaBng nonliving
organic material. But unlike fungi, most
animals do not feed by absorpIon; instead,
animals ingest their food and then use
enzymes to digest it within their bodies.

Cell Structure and SpecializaBon


Animals are eukaryotes, and
like plants and most fungi (but
unlike most proIsts), animals
are mulBcellular. In contrast
to plants and fungi, however,
animals lack the structural
support of cell walls. Instead,
animal cells are held together
by structural proteins, the
most abundant being collagen
(see Figure 6.30), which is
found only in animals.

Denicin de Animal

14

19/8/13

Denicin de Animal
Cell Structure and SpecializaBon

ReproducBon and Development

Many animals have two types of


specialized cells not seen in other
mulIcellular organisms: muscle
cells and nerve cells. In most
animals, these cells are organized
into muscle Issue and nervous
Issue, respecIvely, and are
responsible for moving the body
and conducBng nerve impulses.
The ability to move and conduct
nerve impulses underlies many of
the adaptaBons that dierenBate
animals from plants and fungi,
making muscle and nerve cells
central to what it means to be an
animal.

Most animals reproduce sexually, and the


diploid stage usually dominates the life
cycle. In most species, a small, agellated
sperm ferIlizes a larger, nonmoIle egg,
forming a diploid zygote.
The zygote then undergoes cleavage, a
succession of mitoIc cell divisions without
cell growth between division cycles.
During the development of most animals,
cleavage leads to the formaIon of a
mulIcellular stage called a blastula, which in
many animals takes the form of a hollow ball
(Figure 32.2).
Following the blastula stage is the process of
gastrulaBon, during which layers of
embryonic Issues that will develop into adult
body parts are produced. The resulIng
developmental stage is called a gastrula.

Denicin de Animal

Denicin de Animal

15

19/8/13

Denicin de Animal

Denicin de Animal

AB

CD

apical cell
prostomial ectoderm

112

2a1

1a

{ 1a

{2a2a

2a12

{2a2a

1a1221

prostomial ectoderm
accessory trochoblast
primary trochoblast

1222

111

ectoderm
secondary trochoblast

112

121

secondary trochoblast

122

ectoderm

2a2

ectoderm

{ {
3a1

3a

3a2

2A

apical ganglion

prostomial ectoderm

2a11

122

1a2

1A

ectoderm

3a21
3a22

stomodaeum
3a221 stomodaeum

{3a

222

ectomesoderm
endoderm

3A

{{
C

111

1a121

1a12

2a

{{

{{
1a

{1a1a

{ {
1a11

1a1

as A, except that the right cerebral ganglion develops from 1c112,


there is no ectomesoderm from the B-quadrant, and that the
ectomesoderm originates from 3c21 in the C-quadrant

1d

1D

1d1

{1d1d

apical cell

111

{ {

1d11

112

prostomial ectoderm

1d12

{ {{

primary trochoblast

2d1

2d11
2d12

2d

2d2

2D

apical ganglion
cerebral ganglia

prostomial ectoderm

1d2

{2d2d

21
22

3d
3D

{ 2d2d

111

ectoderm

112

somatoblast
ectoderm, telotroch

circumblastoporal
nerve cord, incl.
posterior part of
brain

ectoderm, telotroch
ectoderm, proctodaeum, ectomesoderm
ectoderm, proctodaeum, ectomesoderm

{4D
4d

mesoderm, endoderm
endoderm

Plate 4. Generalized cell lineage of an annelid. (Based on Podarke and Arenicola; Child 1900; Treadwell 1901). See Figure 22.9,
page 98.

16

19/8/13

Denicin de Animal
ReproducBon and Development

Some animals, including humans,


develop directly into adults through
transient stages of maturaIon, but the
life cycles of many animals also include
at least one larval stage.
A larva is a sexually immature form of
an animal that is morphologically
disBnct from the adult, usually eats
dierent food, and may even have a
dierent habitat than the adult, as in
the case of the aquaIc larva of a
mosquito or a dragony.
Animal larvae eventually undergo
metamorphosis, a developmental
transformaBon that turns the animal
into a juvenile, which resembles an
adult but is not yet sexually mature.

Eubilateria
4
5

ReproducBon and Development

Although adult animals vary widely in


morphology, the underlying geneBc
network that controls animal
development is similar across a broad
range of taxa. All eukaryotes have
genes that regulate the expression of
other genes, and many of these
regulatory genes contain common sets
of DNA sequences called homeoboxes
(see Chapter 21).
Animals share a unique homeobox-
containing family of genes, known as
Hox genes. Hox genes play important
roles in the development of animal
embryos, controlling the expression of
dozens or even hundreds of other
genes that inuence animal
morphology (see Chapter 25).

Deuterostomia
6
7
8

10

Chordata
11 12 13 14

15

Branchiostoma
Mus
Ambulacraria
9/10
11/13 abc

Saccoglossus
Strongylocentrotus

lab

pb Hox3 Dfd Scr

Protostomia
Lox5 Antp

Ecdysozoa
Ubx abd-A abd-B

Tribolium
Priapulus
Spiralia
Lox2 Lox4 Post1 Post2

Capitella
Pecten
Platyhelminthes

Lingula
Xenoturbella
Symsagittifera
Nematostella
Plate 2. Hox genes of a cnidarian and selected representatives of the major bilaterian groups, demonstrating the existence of
series of Hox signatures (see the text). The Eubilaterian Hox signature consists of a group 3 gene and the two anteriormost
central genes. Genes belonging to the same orthology group have the same colour: anterior genes, red; group three, orange;
central group, green; and posterior group, blue. Duplications are not shown. Diagram based on: Branchiostoma: Holland et al.
(2008), Mus: Carroll et al. (2005), Saccoglossus: Aronowicz and Lowe (2006), Strongylocentrotus: Cameron et al. (2006), Tribolium:
Shippy et al. (2008), Priapulus: de Rosa et al. (1999), Capitella: Frbius et al. (2008), Pecten: Canapa et al. (2005), Platyhelminthes:
Olson (2008); Badets and Verneau (2009), Lingula: de Rosa et al. (1999), Xenoturbella: Fritsch et al. (2008), Symsagittifera: Moreno
et al. (2009), Nematostella: Ryan et al. (2007). The platyhelminth sequences are rather divergent so the genes have been inserted
with question marks. See Figure 21.3, page 83.

Denicin de Animal
ReproducBon and Development

Sponges, which are among the simplest


extant animals, have Hox genes that regulate
formaIon of water channels in the body wall,
a key feature of sponge morphology (see
Chapter 33).
In the ancestors of more complex animals,
the Hox gene family underwent further
duplicaIons, yielding a more versaIle
"toolkit" for regulaIng development. In
vertebrates, insects, and most other animals,
Hox genes regulate paLerning of the
anterior-posterior (front-to-back) axis, as well
as other aspects of development. The same
conserved geneIc network governs the
development of both a y and a human,
despite their obvious dierences and
hundreds of millions of years of divergent
evoluIon.

17

19/8/13

The history of animals spans


more than half a billion years

The animal kingdom includes not only


the great diversity of living species,
but also the even greater diversity of
exBnct ones. (Some paleontologists
have esImated that 99% of all animal
species are exInct.)
Various studies suggest that this great
diversity has its origins in
evoluIonary changes that occurred
during the last billion years.
For example, some esImates based
on molecular clocks suggest that the
ancestors of animals diverged from
the ancestors of fungi about a billion
years ago. Similar studies suggest
that the common ancestor of living
animals may have lived someBme
between 675 and 875 million years
ago.

To learn what this common ancestor


may have been like, scienIsts have
sought to idenIfy proIst groups that
are closely related to animals. As
shown in Figure 32.3, a combinaIon
of morphological and molecular
evidence indicates that
choanoagellates are among the
closest living relaBves of animals.
Based on such results, researchers
hypothesize that the common
ancestor of living animals may have
been a staIonary suspension feeder,
similar to present-day
choanoagellates. In this secIon, we
will survey the fossil evidence for
how animals evolved from their
distant common ancestor over four
geologic eras (see Table 25.1 to

Neoproterozoic Era
(1 Billion (1000 mill)-542 Million Years
Ago)
Despite the molecular data indicaIng an earlier origin of

animals, the rst generally accepted macroscopic fossils of


animals range in age from 565 to 550 million years old.
These fossils are members of an early group of mulIcellular
eukaryotes, known collecIvely as the Ediacaran biota.
These soy bodied organisms were named for the Ediacara
Hills of Australia, where they were rst discovered (Figure
32,4).
Similar fossils have since been found on other conInents.
Some are sponges, while others may be related to living
cnidarians. SBll others of these fossil organisms have
proved dicult to classify, as they do not seem to be
closely related to any living animal or plant groups.

Neoproterozoic Era
(1 Billion (1000 mill)-542 Million Years
Ago)
In addiIon to these macroscopic fossils, Neoproterozoic
rocks have also yielded what may be microscopic signs of
early animals. As you read in Chapter 25, 575-million-year-
old microfossils discovered in China appear to exhibit the
basic structural organizaBon of present-day animal
embryos.
However, debate conInues about whether the fossil
embryos are animals or are members of exBnct groups
that are closely related to animals (but are not actually
animals). Though older fossils of animals may be
discovered in the future, the fossil record as it is known
today strongly suggests that the end of the Neoproterozoic
era was a Ime of increasing animal diversity.

18

19/8/13

Paleozoic Era (542-251 Million Years


Ago)
Animal diversicaIon appears to have accelerated dramaIcally
from 535 to 525 million years ago, during the Cambrian period of
the Paleozoic era-a phenomenon o]en referred to as the Cambrian
explosion (see Chapter 25). In strata formed before the Cambrian
explosion, only a few animal phyla can be recognized.
But in strata that are 535 to 525 million years old, paleontologists
have found the oldest fossils of about half of all extant animal
phyla, including the rst arthropods, chordates, and echinoderms.
Many of these disIncIve fossils-which include the rst animals with
hard mineralized skeletons-look quite dierent from most living
animals (Figure 32.5).
But for the most part, paleontologists have established that these
Cambrian fossils are members of extant animal phyla-or at least are
close relaIves.

Paleozoic Era (542-251 Million Years


Ago)
What caused the Cambrian explosion? There are several current
hypotheses. Some evidence suggests that new predator-prey
relaBonships that emerged in the Cambrian period generated
diversity through natural selecBon. Predators acquired novel
adaptaIons, such as forms of locomoIon that helped them catch
prey, while prey species acquired new defenses, such as protecIve
shells.
Another hypothesis focuses on a rise in atmospheric oxygen that
preceded the Cambrian explosion. Greater oxygen availability
would have provided opportuniBes for animals with higher
metabolic rates and larger body sizes to thrive.
A third hypothesis proposes that the evoluBon of the Hox gene
complex provided the developmental exibility that resulted in
variaBons in morphology. These hypotheses are not mutually
exclusive, however; predator-prey relaIonships, atmospheric
changes, and developmental exibility may each have played a role.

Paleozoic Era (542-251 Million Years


Ago)
The Cambrian period was followed by the Ordovician, Silurian, and
Devonian periods, when animal diversity conInued to increase,
although punctuated by episodes of mass exBncBons (see Figure
25.14).
Vertebrates (shes) emerged as the top predators of the marine
food web. By 460 million years ago, groups that diversied during
the Cambrian period were making an impact on land.
Arthropods began to adapt to terrestrial habitats, as indicated by
the appearance of millipedes and cenBpedes.
Fern galls-enlarged caviIes that resident insects sImulate fern
plants to form, providing protecIon for the insects-date back at
least 302 million years, suggesIng that insects and plants were
inuencing each other's evoluBon by that Bme.

19

19/8/13

Paleozoic Era (542-251 Million Years


Ago)
Vertebrates made the transiBon to land
around 360 million years ago and diversied
into numerous terrestrial groups.
Two of these survive today: the amphibians
(such as frogs and salamanders) and the
amniotes (repIles and mammals).
We will explore these groups, known
collecIvely as the tetrapods, in more detail.

NO HAY EXTINCION!!!!

* = caracteres plesiomrcos

EXTINCION EN
REGISTRO FOSIL !!!!

Mesozoic Era (251- 65.5 Million Years


Ago)
No fundamentally new animal groups emerged during the
Mesozoic era.
But the animal phyla that had evolved during the Paleozoic now
began to spread Into new ecological habitats. In the oceans, the
rst coral reefs formed, providing other animals with new marine
habitats. Some repBles returned to the water and succeeded as
large aquaIc predators. On land, descent with modicaBon in
some tetrapods led to the origin of wings and other ight
equipment in pterosaurs and birds. Large and small dinosaurs
emerged, both as predators and herbivores. At the same Ime, the
rst mammals-Bny nocturnal insect eaters- appeared on the
scene. In addiIon, as you read in Chapter 30, owering plants
(angiosperms) and insects both underwent dramaBc
diversicaBons during the late Mesozoic.

20

19/8/13

Cenozoic Era
(65.5 Million Years Ago to the Present)
Mass exBncBons of both terrestrial and marine animals
ushered in a new era, the Cenozoic.
Among the groups of species that disappeared were the
large, nonying dinosaurs and the marine repBles.
The fossil record of the early Cenozoic documents the rise
of large mammalian herbivores and predators as mammals
began to exploit the vacated ecological niches.
The global climate gradually cooled throughout the
Cenozoic, triggering signicant shiys in many animal
lineages. Among primates, for example, some species in
Africa adapted to the open woodlands and savannas that
replaced the former dense forests. The ancestors of our
own species were among those grassland apes.

Animals can be characterized by


"body plans"
Although animal species vary
tremendously in morphology,
their great diversity in form can
be categorized into a relaIvely
small number of major "body
plans.
A body plan is a set of
morphological and
developmental traits, integrated
into a funcIonal whole-the living
animal. (Note that the term plan
here is not meant to imply that
animal forms are the result of
conscious planning or invenIon.)

Like all features of organisms,


animal body plans have evolved
over Ime. Some of the
evoluIonary changes appear to
have occurred early in the history
ofanimal life. For example, recent
research suggests that a key step
in the molecular control of
gastrulaIon has remained
unchanged for more than 500
million years (Figure 32.6). This
early evoluBonary innovaBon
was of fundamental importance:
GastrulaIon explains why most

Animals can be characterized by


"body plans"

animals are not a hollow ball of cells. But


other aspects of animal body plans may have
changed many Imes as various animal
lineages evolved and diversied. As you
explore the major features of animal body
plans, bear in mind that similar body forms
may have evolved independently in two
dierent lineages.
Consider, for example, the group of
invertebrate animals called gastropods (class
Gastropoda). This group includes many
species that lack shells and are referred to as
slugs, along with many shelled species, such
as snails. All slugs have a similar body plan
and hence belong to the same grade (a
group whose members share key biological
features). However, phylogeneIc studies
show that several gastropod lineages
independently lost their shells and became
slugs. As illustrated by this example, a grade
is not necessarily equivalent to a clade (a
group that includes an ancestral species and
all of its descendants).

Un clado ms bien es un grupo polilIco


agrupado por plesiomoras.

21

19/8/13

Animals can be characterized by


"body plans"
Symmetry
One very basic way that animals
can be categorized is by the type of
symmetry of their bodies-or the
absence of symmetry. Most
sponges, for example, lack
symmetry altogether. Among the
animals that do have symmetrical
bodies, symmetry can take dierent
forms. Some animals exhibit radial
symmetry, the form found in a
owerpot (Figure 32.7a). Sea
anemones, for example, have a top
side (where the mouth is located)
and a boLom side.
But they have no front and back
ends and no ley and right sides.

The two-sided symmetry


seen in a shovel is an
example of bilateral
symmetry (Figure 32.7b). A
bilateral animal has two
axes of orientaIon: front to
back and top to boLom.

Animals can be characterized by


"body plans"
Symmetry
Thus, such animals have a dorsal
(top) side and a ventral (boLom)
side, as well as a ley side and a
right side and an anterior (front)
end with a mouth and a posterior
(back) end. Many animals with a
bilaterally symmetrical body plan
(such as arthropods and
mammals) have sensory
equipment concentrated at their
anterior end, including a central
nervous system ("brain") in the
head-an evoluIonary trend called
cephalizaIon (from the Greek
kephale, head).

The symmetry of an animal generally


ts its lifestyle. Many radial animals are
sessile (living aLached to a substrate) or
planktonic (driying or weakly
swimming, such as jellysh, more
accurately called jellies). Their
symmetry equips them to meet the
environment equally well from all sides.
In contrast, bilateral animals typically
move acIvely from place to place.
Most bilateral animals have a central
nervous system that enables them to
coordinate the complex movements
involved in crawling, burrowing, ying,
or swimming. Fossil evidence indicates
that these two fundamentally dierent
kinds of symmetry have been present
for at least 550 million years.

Animals can be characterized by


"body plans"
Tissues

Animal body plans also vary according to the


organizaIon of the animal's Issues. True
Issues are collecIons of specialized cells
isolated from other Issues by membranous
layers. Sponges and a few other groups lack
true Issues. In all other animals, the embryo
becomes layered through the process of
gastrulaIon (see Figure 32.2). As
development progresses, these concentric
layers, called germ layers, form the various
Issues and organs of the body.
Ectoderm, the germ layer covering the
surface of the embryo, gives rise to the outer
covering of the animal and, in some phyla, to
tlle central nervous system.
Endoderm, the innermost germ layer, lines
the developing digesIve tube, or
archenteron, and gives rise to the lining of
the digesIve tract (or cavity) and organs such
as the liver and lungs ofvertebrates.

Animals that have only these two germ layers are


said to be diploblasIc. Diploblasts include the
animals called cnidarians, jellies and corals, for
example) as well as the comb jellies (see Chapter
33).

All bilaterally symmetrical animals have a third germ


layer, called the mesoderm, betweeen the ectoderm
and endoderm. Thus, animals with bilateral
symmetry are also said to be triploblasIc (having
three germ layers).

In triploblasts,the mesoderm forms the muscles and


most other organs between the digesIve tract and
the outer covering of the animal.

Triploblasts include a broad range of animals, from


atworms to arthropods to vertebrates. (Although
some diploblasts actually do have a third germ layer,
it is not nearly as well developed as the mesoderm
of animals considered to be triploblasIc.)

22

19/8/13

Animals can be characterized by


"body plans"
Body CaviBes
Most triploblasIc animals possess a
body cavity, a uid- or air lled
space separaIng the digesIve tract
from the outer body wall. This body
cavity is also known as a coelom
(from the Greek koilos, hollow).
Also-called "true" coelom forms
from Issue derived
from mesoderm. The inner and
outer layers of Issue that surround
the cavity connect dorsally and
ventrally and form
structures that suspend the internal
organs. Animals that possess a true
coelom are known as coelomates

Some triploblasIc animals have a


body cavity that is formed from
mesoderm and endoderm (Figure
32,8b). Such a cavity is called a
"pseudocoelom" (from the Greek
pseudo, false), and animals that
have one are pseudocoelomates.
Despite its name, however, a
pseudocoelom is not false; it is a
fully funcIonal body cavity.
Finally, some triplobasIc animals
lack a body cavity altogether
(Figure 32.8c). They are known
collecIvely as acoelomates (from
the Greek a, without).

Animals can be characterized by


"body plans"
Body CaviBes
Abody cavity has many funcIons.
Its uid cushions the suspended
organs, helping to prevent internal
injury. In soy-bodied coelomates,
such as earthworms, the coelom
contains noncompressible uid that
acts like a skeleton against which
muscles can work. The cavity also
enables the internal organs to grow
and move independently oyhe
outer body wall. It were not for
your coelom, for example, every
beat of your heart or ripple of your
intesIne would warp your body's
surface.

Current phylogeneIc
research suggests that true
coeloms and
pseudocoeloms have been
gained or lost mulIple
Imes in the course of
animal evoluIon. Thus, the
terms coelomates and
pseudocoelomates refer to
grades, not clades.

Protostome and Deuterostome


Development
Based on certain aspects of early
development, many animals can be
categorized as having one of two
developmental modes:
protostome development or deuterostome
development.
These modes can generally be disInguished
by dierences in cleavage, coelom formaIon,
and fate of the blastopore.

23

19/8/13

Protostome and Deuterostome


Development
Cleavage

Cleavage

A paLern in many animals with


protostome development is spiral
cleavage, in which the planes ofcell
division are diagonal to the verIcal axis
oyhe embryo; as seen in the eight-cell
stage, smaller cells lie in the grooves
between larger, underlying cells (Figure
32,9a, ley).
Furthermore, the so-called determinate
cleavage of so meanimals with
protostome development rigidly casts
("determines~) the developmental fate
of each embryonic cell very early. A cell
isolated at the four-cell stage from a
snail, for example, ayer repeated
divisions will form an inviable embryo
that lacks many parts.

mosaico

In contrast to the spiral cleavage paLern, deuterostome


development
is predominantly characterized by radial cleavage. The
cleavage planes are either parallel or perpendicular to the
verIcal
axis of the embryo; as seen in the eight-cell stage, the Iers
ofcells
are aligned, one directly above the other (see Figure 32.9a,
right).
Most animals with deuterostome development also have
indeterminate cleavage, meaning that each cell produced by
early cleavage divisions retains the capacity to develop into
acorn
plete embryo. For example, if the cells of a sea urchin embryo
are
isolated at the four-cell stage, each will form acomplete larva.
It is
the indeterminate cleavage oyhe human zygote t1lat makes
idenIcal
twins possible. nlis characterisIc also explains the
developmental
versaIlity of embryonic stem cells that may provide new
ways to overcome a variety ofdiseases (see Chapter 20)

regulatorio

Protostome and Deuterostome


Development
Coelom Forma2on

Fate of the Blastopore

Another dierence betv:een


protostome and deuterostome
development is apparent later in
the process. During gastrulaIon, an
embryo's developing digesIve tube
iniIally forms as a blind pouch, the
archenteron, which becomes the
gut (Figure 32.9b).
As the archenteron forms in
protostome development, iniIally
solid masses of mesoderm split and
form the coelom. In contrast, in
deuterostome development, the
mesoderm buds from the wall
oyhe archenteron, and its cavity
becomes the coelom.

Protostome and deuterostome development


oyen dier in the fate oyhe blastopore, the
indentaIon that during gastrulaIon leads to
the formaIon of the archenteron (Figure
32.9c). Ayer the archenteron develops, in
most animals a second opening forms at the
opposite end of the gastrula. UlImately, the
blastopore and this second opening become
the two openings of the digesIve tube (the
mouth and the anus). In protostome
development, the mouth generally develops
from the rst opening, the blastopore, and it
is for this characterisIc that the term
protostome derives (from the Greek protos,
rst, and stoma, mouth).
In deuterostome (from the Greek deuteros,
second) development, the mouth is derived
from the secondary opening, and the
blastopore usually forms the anus.

24

19/8/13

New views of Animal phylogeny are emerging from


molecular data

Zoologists currently recognize about three


dozen animal phyla. But the relaIonships
between these phyla conInue to be debated.
Although it might be frustraIng that the
phylogenies in textbooks cannot be
memorized as set-in-stone truths, the
uncertainty inherent in these diagrams is a
healthy reminder that science is an ongoing,
dynamic process of inquiry.

New views of Animal phylogeny are emerging from


molecular data

Researchers have long based their hypotheses


about animal phylogeny on morphological
data. In the late 1980s, biologists also began
to study the molecular systemaIcs of animals.
AddiIonal clues about animal phylogeny have
come from new studies of lesser-known phyla,
along with fossil analyses that can help clarify
which traits are ancestral in various animal
groups and which are derived.

New views of Animal phylogeny are emerging from


molecular data

As you read in Chapter 26, phylogeneIc


systemaIsts seek to classify organisms based on
common descent. Their goal is to place organisms
into groups that correspond to clades, each of
which includes an ancestral species and all of its
descendants. Based on cladisIc methods, a
phylogeneIc tree takes shape as a hierarchy of
clades nested within larger clades-the ner
branches and the thicker branches of the tree,
respecIvely.

25

19/8/13

New views of Animal phylogeny are emerging from


molecular data
Clades are inferred from shared derived characters that
are unique to members oyhe clade and their common
ancestor.
For example, a clade might be inferred
from key anatomical and embryological
similariIes that researchers conclude
are homologous. In recent years, molecular
data such as DNA sequences
have provided another source of informaIon
for inferring common ancestry
and dening clades.

New views of Animal phylogeny are emerging from


molecular data

But whether the


data used are "tradiIonal" morphological
characters or "new" molecular sequences
or a combinaIon of both, the
overall goal is the same-to reconstruct
the evoluIonary history of life. To get a sense of the
debates that occur
in animal systemaIcs, we'll compare
a tradiIonal view of animal phylogeny,
based primarily on morphological data
(gure 32.10), with a view based mainly
on molecular data (gure 32.11).

New views of Animal phylogeny are emerging from


molecular data - Points of Agreement

The morphological and molecular trees agree


on a number of
major features of animal phylogeny. Ayer
reading each point,
see how the statement is reected in the
phylogeneIc trees in
Figures 32.10 and 32.1l.

26

19/8/13

Espiral o radial: opImice

S
R

Espiral o radial: no son sinapomoras

I
R

S
S
R
S
S

R
S

27

19/8/13
Higher-level metazoan relationships: recent progress

Consequently, myriad phylogenetic hypotheses have


been published in the past decade alone, and even the most
dedicated systematic researcher may from time to time fail
to see the consensus in this deep forest of phylogenetic
trees. In the present contribution we review some of the
most recent phylogenetic analyses of higher-level metazoan
relationships, with emphasis on the newest developments,
mostly based on phylogenomic studies. By pointing out

congruence and discrepancies between various hypotheses


we explore the current consensus in metazoan phylogeny
(summarized in Fig. 1), and discuss our present knowledge
in relation to morphological and developmental character
evolution. We also appraise the merits and limitations of
several alternative molecular phylogenetic approaches that
are currently used to address the vexing question of
metazoan interrelationships.

Fig. 1 Summary of relationships


within Metazoa. Nodes labelled
with circled letters (taxon-name
abbreviations) have received
broad consensus

Aco
ela

Deutero Ecdysozoa
stomia

Lophotrochozoa

Lophotrochozoa comprise two groups,


the trochozoans and the lophophorata
Lophotrochozoa
Larvas trocforas en
general
lophos = indicaIng a
crested or tuyed part
trocho = wheel
zoon = animals
crest/wheel animals

New views of Animal phylogeny are emerging from


molecular data - Points of Agreement

I. All animals share a common


ancestor.
Both trees indicate that the animal
kingdom is monophyleIc,
represenIng a dade called Metazoa.
In other words, if we could trace all
extant and exInct animal lineages
back to their origin, they would
converge on a common ancestor.
2. Sponges are basal animals. Among
the extant taxa, sponges (formerly,
the phylum Porifera) branch from the
base of both animal trees. Some
molecular analyses suggest that
sponges are paraphyleIc, falling into
at least the two phyla shown in Figure
32.11.

3. Eumetazoa is a dade of animals


with true Issues.
All animals except for sponges and a
few other groups belong to a clade
ofeumetazoans ("true animals"). True
Issues evolved in the common
ancestor of living eumetazoans.
Basal members of the eumetazoan
clade include the phyla Ctenophora
(comb jellies) and Cnidaria.
These basal eumetazoans are
diploblasIc and generally feature
radial symmetry.

28

19/8/13

New views of Animal phylogeny are emerging from


molecular data - Points of Agreement
4. Most animal phyla belong to 5. Chordates and some other
the dade Bilateria.
phyla belong to the
Bilateral symmetry and the
clade Deuterostomia.
presence of three germ layers The term deuterostome refers
are shared derived characters
not only to a mode of animal
that help to dene the clade
development, but also to the
Bilateria.
members of a dade that
includes vertebrates and other
This clade contains the
majority of animal phyla, and
chordates. (Note, however,
its members are known as
that the tradiIonal and
biJatcrians.
molecular views of animal
phylogeny disagree as to
The Cambrian explosion was
which other phyla are also
primarily a rapid diversicaIon
deuterostomes.)
of the bilaterians.

FIN

29