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
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
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
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
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
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
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
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
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
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).
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
19/8/13
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.)
Origen de Metazoa
Teora
colonial-
Haeckel
Teora
celularizacin
(sincicial)-Hadzi,
Steinbck
19/8/13
Volvox
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].
Teora colonial-Haeckel
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.
1 Animal Evolution
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
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
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
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
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.
mitosis
water current
nerve
germ cells
basement
membrane
sensory cell
cadherins
occluding junction
circumblastoporal
nerve
Ancestral sponge
Primitive sponge
Neurogastraea
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
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
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.
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 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
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
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
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
Denicin de Animal
14
19/8/13
Denicin
de
Animal
Cell
Structure
and
SpecializaBon
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
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
Eubilateria
4
5
Deuterostomia
6
7
8
10
Chordata
11 12 13 14
15
Branchiostoma
Mus
Ambulacraria
9/10
11/13 abc
Saccoglossus
Strongylocentrotus
lab
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
17
19/8/13
Neoproterozoic
Era
(1
Billion
(1000
mill)-542
Million
Years
Ago)
Despite
the
molecular
data
indicaIng
an
earlier
origin
of
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
19
19/8/13
NO HAY EXTINCION!!!!
* = caracteres plesiomrcos
EXTINCION
EN
REGISTRO
FOSIL
!!!!
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.
21
19/8/13
22
19/8/13
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.
23
19/8/13
Cleavage
mosaico
regulatorio
24
19/8/13
25
19/8/13
26
19/8/13
S
R
I
R
S
S
R
S
S
R
S
27
19/8/13
Higher-level metazoan relationships: recent progress
Aco
ela
Deutero Ecdysozoa
stomia
Lophotrochozoa
28
19/8/13
FIN
29