CONSIDERACIONES ACERCA DEL ORIGEN Y

EVOLUCIÓN DE LOS CORDADOS

PAOLA ESCOBAR RAMOS

BIÓLOGO, DOCENTE
MSC. CIENCIAS BIOLÓGICAS
Humanos modernos
Primeros homínidos

Mamíferos modernos
Primeras ballenas

Primeras plantas con flor

Primeros pájaros
Primeros Dinosaurios y mamíferos
Primeros reptiles

Primeros anfibios terrestres

Primeros insectos
Primeras plantas terrestres
Primeros peces
Primeros cordados
Primeras formas de vida pluricelulares
Primeras formas de vida unicelulares
PHYLLUM CHORDATA: Aportes biológicos

• Endoesqueleto, vértebras

• Crecimiento

• Faringe perforada

• Origen de branquias y mandíbula

• Hábitos depredadores y cerebros altamente

desarrollados

• Apéndices pareados
 CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN
• Aparición, período Cámbrico 530 millones de años
PHYLUM CHORDATA: Notochord
• Primeros cordados, cuerpo suave
In a lancelet, the simplest chordate, the
flexible notochord persists throughout
life and aids swimming by giving muscles

Cinco carácteres diagnósticos:

something to pull against. In the lancelet
• these muscles form a series of discrete
blocks that can easily be seen. More
advanced chordates have jointed appendages.

Notochord

1. Notocordio
Water
Lancelets lack pigment in their
Dorsal nerve skins, and so are transparent.
cord

Oral hood
with tentacles

2. Cordon nervioso dorsal Gill slits

in pharynx

Lancelets are filter-feeders

with highly reduced Atrium

3. Hendiduras faringeas
sensory systems. The
Atriopore
animal has no head, eyes,
ears, or nose. Instead,
sensory cells that detect
chemicals line the oral Anus
Intestine
tentacles.

Lancelets feed on microscopic

4. Endostilo o glándula tiroide protists caught by filtering them

through cilia and gills on the
pharyngeal slits. As the cilia that
line the front end of the
Unlike that of vertebrates, the
skin of a lancelet has only a
gut passage beat, they draw water single layer of cells.
through the mouth, through the
pharynx, and out the slits.

5. Cola post anal

FIGURE 48.4
Evolution of a notochord. Vertebrates, tunicates, and lancelets are chordates (phylum Chordata), coelomate animals with a flexible rod,
the notochord, that provides resistance to muscle contraction and permits rapid lateral body movements. Chordates also possess
pharyngeal slits (reflecting their aquatic ancestry and present habitat in some) and a hollow dorsal nerve cord. In vertebrates, the
 CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN
48.2 Nonvertebrate chordates have a notochord but no backbone.

The Nonvertebrate Chordates The tadpolelike larvae of tunicates plainly exhibit all
the basic characteristics of chordates and mark the tun
• Endoesqueleto; locomocion, soporte y protección
Tunicates
The tunicates (subphylum Urochordata) are a group of
cates as having the most primitive combination of featur
found in any chordate (figure 48.5c). The larvae do n
mals. In colonial tunicates, there may feed and have a poorly developed gut. They remain fre
about 1250 species of marine animals. Most of them are
be a common sac and a common swimming for only a few days before settling to the bo
sessile as adults (figure 48.5a,b), with only the larvae hav-
opening to the outside. There is a tom and attaching themselves to a suitable substrate b
ing a notochord and nerve cord. As adults, they exhibit

Oseo o cartilaginoso, bajo la piel

group of Urochordates, the Larvacea, means of a sucker.
• neither a major body cavity nor visible signs of segmenta-
which retains the tail and notochord
tion. Most species occur in shallow waters, but some are
Tunicates change so much as they mature and adju
into adulthood. One theory of verte- developmentally to a sessile, filter-feeding existence th
found at great depths. In some tunicates, adults are colo-
brate origins involves a larval form, it would be difficult to discern their evolutionary rel
nial, living in masses on the ocean floor. The pharynx is
perhaps that of a tunicate, which ac- tionships by examining an adult. Many adult tunicates s
lined with numerous cilia, and the animals obtain their
quires the ability to reproduce. crete a tunic, a tough sac composed mainly of cellulos
food by ciliary action. The cilia beat, drawing a stream of
• Terrestres Lancelets
water into the pharynx, where microscopic food particles
are trapped in a mucous sheet secreted from a structure
The tunic surrounds the animal and gives the subphylu
its name. Cellulose is a substance frequently found in th
cell walls of plants and algae but is rarely found in an
called an endostyle.
Lancelets are scaleless, fishlike marine
chordates a few centimeters long that
occur widely in shallow water
• Con mandíbulas throughout the oceans of the world.
Lancelets (subphylum Cephalochor-
data) were given their English name Hypophyseal duct Incurrent siphon
because they resemble a lancet—a
small, two-edged surgical knife. There Nerve ganglion
Pharynx

Sin esqueleto óseo o cartilaginoso

are about 23 species of this subphy-
• lum. Most of them belong to the
genus Branchiostoma, formerly called FIGURE 48.6
Excurrent
siphon
Endostyl
Amphioxus, a name still used widely. In Lancelets. Two lancelets, Branchiostoma lanceolatum (phylum Chordata, subphylum
lancelets, the notochord runs the en- Cephalochordata), partly buried in shell gravel, with their anterior ends protruding. The Gill slit
tire length of the dorsal nerve cord muscle segments are clearly visible; the square objects along the side of the body are
Intestine

Marinos
gonads, indicating that these are maleGenital
lancelets. Tunic
and persists throughout the animal’s
•
duct
life.
Lancelets spend most of their time Gonad
Stomach
partly buried in sandy or muddy substrates, with Oral hood Notochord Dorsal Heart
with tentacles
only their anterior ends protruding (figure 48.6). nerve cord Intestine
They can swim, although they rarely do so. Their (b)

• Suspensívoros muscles can easily be seen as a series of discrete

blocks. Lancelets have many more pharyngeal gill
slits than fishes, which they resemble in overall
Mouth
(incurrent siphon)
Atriopore
(excurrent siphon)
AnusDorsal nerve co
shape. They lack pigment in their skin, which has
only a single layer of cells, unlike the multilayered Gill slits Gonad Atrium Atriopore
in pharynx
skin of vertebrates. The lancelet body is pointed at
• Colágeno both ends. There is no distinguishable head or
sensory structures other than pigmented light re-
FIGURE 48.7
The structure of a lancelet.
Pharynx
slits diagram showsStomach
with gillThis the path through whichNotochord
ceptors. the lancelet’s cilia pull
(c) water. Heart
Lancelets feed on microscopic plankton, using a
current created by beating cilia that lines the oral FIGURE 48.5
hood, pharynx, and gill slits (figure 48.7). The gill Tunicates (phylum Chordata, subphylum Urochordata).
slits provide an exit for the water and are an adaptation for group of(a) chordates.
The sea peach, The Halocynthia auranthium.
recent discovery of(b)fossil
Diagram
formsof the
filter feeding. The oral hood projects beyond the mouth similar structure
to livingoflancelets
an adult tunicate.
in rocks (c)
550Diagram
millionof years
the structure
old— of a
 Trunk Collar Proboscis

Branchial
pouch Proboscis
Branchial coelom
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60
Mouth Stomochord
Trunk Collar
coelom coelom

Anus Pharyngeal slit in pharynx

HEMICHORDATA
FIGURE 2.8 Hemichordate, generalized acorn worm. Proboscis, collar, and trunk regions are shown in partial cutaway view,
revealing the coelom in each region and the associated internal anatomy of the worm. Within the proboscis is the stomochord, an
Tentacles
extension of the digestive tract. The food-laden cord of mucus
Collar(spiral arrow at right) enters the mouth together with water. Food is
directed through the pharynx into the gut. Excess water exits via the pharyngeal slits. Several slits open into each branchial pouch, a
Proboscis
common compartment with a branchial pore that opens to the outside environment. Arms

Modified from Gutmann.

Anus Mouth

Blastocoel in length, but most are shorter than this. Most live in mucus-
lined burrows and have a body with three regions—proboscis,
Coelomic
Nephridium compartment
Oral collar, trunk—each with its own coelom (figure 2.10a–c).
lamella
Nephridial ductArms
Mouth The proboscis,
Tube
used in both locomotion and feeding, includes
Preoral a muscular outer wall that encloses a fluid-filled coelomic
hood space. Muscular control over the shape of the proboscis gives
Zooid
Stalk
the animal a useful probe to shape a tunnel or inflate itself
Gut
Trunk against the walls of the burrow to anchor its body in place
Nephridial pore (figure 2.10b). Tucked away in their burrows, many species
Pulsatile vesicle
Circumoral ingest loosened sediment, extract the organic material it con-
band
tains, pass the spent sediment through their simple gut, and
Telotroch deposit
(a) (b) a casting (fecal waste) on the surface of the substrate
where changing tides flush it away. Some wide-bodied,
FIGUREFIGURE 2.9 Hemichordate,
2.14 Hemichordata,generalized tornaria (a) The sessiledeep-sea
Pterobranchia. pterobranchenteropneusts crawl
Rhabdopleura. Notice thatand glide along
this pterobranch the abyssal
has the
larva. The same
simple gutplan
body begins
as anatacorn
the mouth under a preoral
worm—proboscis, hood these three
collar, trunk—but oceanfeatures are modified and the whole animal lives in a tube.
bottom.
(a)
Dorsal and tubular
Myomeres nerve cord
Dorsal fin Ocellus
Notochord
Caudal fin

Hindgut Atriopore
Anus Midgut cecum
Ileocolic ring
Esophagus Oral hood
Postanal tail Midgut Pharyngeal slit

(b)
CEPHALOCHORDATA
Ocellus Notochord Wheel organ
Dorsal and Velar tentacle
tubular nerve cord Velum
Muscles Right
Testes stomach
lobe
Incurrent Ovary Ovary
Cecum
Oral Pharynx Testes
siphon (mouth) Intestine Intestine Left
tentacle
stomach
Dorsal lamina Cilated funnel Mouth Trunk
lobe
Right
Endostyle
Ganglion Anus stomach Esophagus
Incurrent lobe
Heart Pharyngeal
Subneural gland Anus slit
Pharyngeal
slit Prebuccal
Atrial Endostyle gland
Excurrent Excurrent
siphon opening Mouth
(atrial) Feeding-
Screen
siphon filter
Tunic Gelatinous
matrix
Atrium (“House”)
Tail
(a)

Dorsal Notochord Notochord

Endostyle lamina

Stigmata

Dorsal
(b)

Anus
(c)

UROCHORDATA
Heart FIGURE 2.26 Urochordata, Larvacea (Appendicularia), Oikopleura albicans. (a) Oikopleura is shown within its gelat
house.The animal’s feeding-filter obtains food from the incurrent of circulating water (small arrows).This larvacean resides at the ba
Gonad
Ciliated band the screen where it sucks food off these screens. (b and c) Enlarged lateral and ventral views, respectively, of the isolated larvacean. T
undulating tail, supported by a notochord, is active in producing the current of food-bearing water that moves through the internal
Stomach
channels of the house and through the feeding-filter.
(a) After Flood; (b, c) after Alldredge.
Stigmata
(b) Vessels of
branchial
new house continues with the addition of feeding screens feeders. These tiny organisms that escape the clutc
basket
and filters. Sometimes within the space of only a few min- other suspension feeders fall prey to the efficient fil
Dorsal lamina utes, the new house is complete and the larvacean is once gear of larvaceans. Larvacea are able to sift through
again feeding actively. volumes of water, ingest a wide range of plankton
All species, except one, are monoecious, and most of including the very tiniest, and proliferate rapidly in res
these are protandrous; that is, sperm and eggs are produced to local blooms in food supply.
Organs
by the same gonad (of the same individual) but at different The trunk of the larvacean holds its major body o
Heart times during its life. Maturation is so rapid that within 24 to although which organs are present varies among the
Posterior 48 hours of fertilization, miniature larvaceans secrete a
Subendostyle families of Larvacea. Members of the smallest famil
vessel vessel house and are set up for feeding. Kowalevskiidae, lack endostyle and heart. In the Fritilla
(d) Their rapid reproduction and special feeding appara- the stomach consists of only a few cells. In the Oikop
 CORDADOS: CARACTERÍSTICAS QUE LOS Kardong: Vertebrates: 2. Origin of Chordates Text

DISTINGUEN - NOTOCORDA O NOTOCORDIO

Comparative Anatomy,
Function, Evolution, Fifth
Edition

• Mesodermo br
Connective tissue m
sheath
ve
• Encima del celoma, debajo del sistema Collagenous on
sheath it
nevioso central fo
Vacuole in fu
notochordal
cell
na
• Compuesta por: m
ic

• Núcleo de células y liquido

(a)
Ph
• Células vacuoladas o entre las células del Without
notochord
With
notochord
A
ur
núcleo de la notocorda im
th
pi
• Propiedades mecánicas= tejido fibroso+núcleo in
all
liquido (b) (c) (d)
us
bu
an
FIGURE 2.5 Notochord. (a) Cross section of the
ha
• Notocorda, órgano hidrostático notochord of a frog tadpole. (b) The notochord lies above the
body cavity and is axially incompressible; that is, it resists br
shortening in length. (c) The notochord is flexible laterally, ar
however. (d) As seen from above, the consequences of muscle tio
contraction in a body with and without a notochord.Without a m
notochord, lateral muscle contraction telescopes the body ro
 ←⎯ Acraniata ⎯→ ⏐ ←⎯⎯⎯⎯⎯⎯
⏐ ←⎯⎯⎯⎯ Ag
CORDADOS: CARACTERÍSTICAS QUE LOS
⏐ ←⎯⎯⎯⎯⎯⎯
DISTINGUEN - CORDÒN NERVIOSO

• Tubo hueco, deriva del ectodermo

Brain Dorsal nerve cord

• Invaginación
56 Kardong: Vertebrates: 2. Origin of Chordates Text © The McGraw−Hill
Comparative Anatomy, Companies, 2009
Function, Evolution, Fifth

• Placa neural, dorsal (células

Edition

nerviosas del embrión)

mucus lining the pharynx snatches food particles from suspen-
Dorsal Tubular Nerve Cord
Annelid or
sion. Sets of cilia, also lining the pharynx, produce the water Chordate
Arthropod
current. Other cilia gather the food-laden mucus and pass it

In most invertebrate phyla that have a

into the esophagus. This mucus and cilia system is especially Dorsal
hollow

Placa neural se enrolla

efficient in small, suspension-feeding organisms, those that
•
nerve cord
extract food floating in water. Such a feeding system is preva-
lent in primitive chordates and in groups that preceded them.nerve cord, it is ventral to the alimen-
In the earliest vertebrates that depended upon gill res-
Notochord

tary canal and is solid, but in chordates

piration to support an active lifestyle, mucus and cilia served
less well. Cilia are weak pumps, ineffective against gill resis-
Digestive tract

Por encima del tubo digestivo y la

Solid nerve cord
tance. In such vertebrates, a pharyngeal pump worked by
• muscles takes the place of cilia to now move the water that the single cord is dorsal to the alimen-
(a) (b)

ventilates the gills. The muscular pump, in place of mucus

notocorda
and cilia, also becomes the basis for procurement and pro- tary canal and is a tube (although the
cessing of large food items. Slits still serve as convenient exit
FIGURE 2.6 Dorsal hollow nerve cord. (a) Basic body
plan of an annelid or arthropod. In such animals, a definitive nerve
cord, when present, is ventral in position, solid, and lies below the
portals for excess or spent water, while adjacent gill struc-
tures function in respiration. In fishes and aquatic amphib- hollow center may be nearly obliter-
digestive tract. (b) Basic chordate body plan.The nerve cord of
chordates lies in a dorsal position above the digestive tract and
ians, the pharyngeal slits that appear during embryonic notochord. Its core is hollow, or more correctly, it has a fluid-
development usually persist into the adult and form the exit ated during growth). The anterior end
filled central canal, the neurocoel, indicated as the white spot in

• Neurocele channel through which water associated with feeding and the dorsal hollow nerve cord.
respiration flows. In vertebrates that reside on land, how-
ever, the embryonic pharyngeal slits normally never open
becomes enlarged to form the brain.
and thus do not give rise to any adult derivative. form the central nervous system do not amass as thickened
Why cilia are replaced by muscles
The hollow cordinstead,iscellsproduced
surface plates (placodes); individually move in the
inward to assemble into the basic nervous system. Further, the
as body size increases (p. 128)
embryo majorby the
nerve cord in mostinfolding ofposition,
invertebrates is ventral in
below the gut, and solid. In chordates, however, the nerve
ectodermal
 CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN - HENDIDURAS FARÍNGEAS
• Faringe parte del tubo digestivo

• La faringe tuvo aberturas longitudinales

• Función respiración

• Alimentación

• Flujo unidireccional del agua

• Corriente de agua participa en el intercambio

gaseoso

• Oxígeno se incorpora a los capilares, el CO2 sale

al agua

CORDADOS: CARACTERÍSTICAS QUE LOS
DISTINGUEN - HENDIDURAS FARÍNGEAS
tebrates, the nerve cord passes through
the protective neural arches of the ver-
tebrae, and the anterior brain is sur-
rounded by a bony or cartilaginous
cranium.

• Cordados primitivos las

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6:44 PM Page 70

hendiduras se expanden, cesta

faringea o branquial Muscles

Incurrent
Oral
Endostyle siphon (mouth)
tentacle
Dorsal lamina Cilated funnel

• Moco pegajoso, capturar las Pharyngeal Pouches

Ganglion

Subneural gland

partículas
Atrial

and Slits
Excurrent
siphon
(atrial)
Ventral siphon
blood Tunic
sinus

Pharyngeal slits are perforated slitlike

Atrium

Dorsal

openings that lead from the pharyn- lamina

Grupo de cilios
(a) Endostyle

• Stigmata

geal cavity to the outside. They are Dorsal

formed by the inpocketing of the out-

Anus
side ectoderm (pharyngeal grooves)
• Persisten en los peces y anfibios Heart

Gonad
Ciliated band
Stomach
Stigmata
(b) Vessels of
branchial
basket
Dorsal lamina

(c) Organs
Heart
Posterior Subendostyle
 tebrates, the nerve
tostomes. Embryological cord passes
characteristics, through
modern molecular
phylogenies, and the fossil record all imply that there was
rigid inte
skin. Th
theanprotective neural
ancient and fundamental arches
divergence betweenof
the the
proto- ver-
port, and

CORDADOS: CARACTERÍSTICAS QUE LOS

stomes and deuterostomes.
tebrae,Chordates
and evolved
the within
anterior brain is
the deuterostomes. Their
mouth forms opposite to the blastopore, their cleavage gen-
are terres
sur-
cles. But
animals,
rounded by a bony or cartilaginous
DISTINGUEN - ENDOSTILO erally is radial, their coelom is an enterocoelom, and their
cranium.
skeleton arises from mesodermal tissues of the embryo. But
we should be clear about the character of the chordates
laginous
involve
and uroc
themselves. It is easy to forget that two of the three chordate sheet of
taxa are technically invertebrates—the Cephalochordata streams o
and the Urochordata. Strictly speaking, the invertebrates All
Pharyngeal
include all animalsslits between
except members ofaortic arches
the vertebrates. share a c
The earliest chordate fossils appear in the Cambrian mental f
period, about 530 million years ago. Although later chor- thyroid g
dates evolved hard bones and well-preserved teeth that left central n
a substantial fossil testimony to their existence, ancestors These fi
to the first chordates likely had soft bodies and left almost together,
no fossil trace of the evolutionary pathway taken from at each c

Gut Notochord

Endostilo predecesor del la

Dorsal hollow
• nerve cord
Pharyngeal Pouches Anus (a)

tiroides, homología and Slits

Atriopore

Postanal tail
Atrium
Pharyngeal slit in pharynx

Pharyngeal slits
(c)
are perforated
(b)
slitlike
openings that Dorsal
leadhollow from the pharyn-
• Metabolismo del yodo geal cavity to the
nerve cord*
outside. They are
Notochord*
formed by the Gut
inpocketing of the out- FIGURE 2
side ectoderm (pharyngeal
Pharynx* grooves)
(a) A single str
pharynx, and t
Atrium
lower chordat
Coelom common enclo
the single atrio
Endostyle*
the floor of th
Body wall showing the tu
(c) Cross sect
(c) (b) indicate chord
 (sharks)
CORDADOS: CARACTERÍSTICAS QUE LOS
⎯⎯⎯⎯ Chordata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
DISTINGUEN - RABO O COLA POSTANAL
⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Vertebrata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Craniata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⏐ ←⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯ Gnathostomata ⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
⏐ ←⎯⎯⎯⎯ Pisces ⎯⎯→ ⏐ ←⎯⎯⎯⎯⎯⎯⎯ Tetrapoda ⎯⎯⎯⎯⎯⎯⎯→
⎯⎯⎯ larvas de tunicados⏐ y←⎯⎯⎯ Amniota ⎯⎯⎯⎯⎯→
Motilidad,⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯⎯→
• Anamniota
anfioxus

• Propulsión,
e evagination, peces
or outpocketing, Segmented myotomes between
endodermal lining of the phar- septa anchored to notochord
• Coxis,
haryngeal humanos
pouches). In aquatic
tes, the two pockets break
h the pharyngeal cavity where
• Rabo, mamíferos
eet to form the pharyngeal slit.
Postanal tail
iotes these pockets may not
• Prolongación
hrough the pharyngealdelcavity
cuerpo
ly grooves are formed instead Postanal Tail
In tetrapod (four-footed) verte- The postanal tail, together with so-
the pharyngeal pouches give matic musculature and the stiffening
 HISTORIA EVOLUTIVA DE LOS CORDADOS

• Debate acerca de la evolución del grupo, XIX

• Sin certeza sobre la apariencia y posición filogenética

dentro de los Metazoarios

• Primeros cordados con hendiduras faríngeas

(hendiduras branquiales)

• Cefalización

• Hendiduras branquiales caracter primitivo,

Deuterostomados
 developmentally to a sessile, filter-feeding existence that
found at great depths. In some tunicates, adults are colo-
it would be difficult to discern their evolutionary rela-
nial, living in masses on the ocean floor. The pharynx is
tionships by examining an adult. Many adult tunicates se-
lined with numerous cilia, and the animals obtain their
crete a tunic, a tough sac composed mainly of cellulose.
food by ciliary action. The cilia beat, drawing a stream of
The tunic surrounds the animal and gives the subphylum
water into the pharynx, where microscopic food particles
its name. Cellulose is a substance frequently found in the
HISTORIA EVOLUTIVA DE LOS CORDADOS
are trapped in a mucous sheet secreted from a structure
called an endostyle.
cell walls of plants and algae but is rarely found in ani-

• Ancestro cordado: Hypophyseal duct Incurrent siphon

Nerve ganglion
Pharynx

Organismo filtrador
Excurrent
• siphon
Endostyle

Gill slit
Intestine
• Forma de tunicado Genital duct
Tunic

Gonad
Stomach
Heart

• Larva en forma de renacuajo (b)

Mouth Atriopore
(incurrent siphon) (excurrent siphon)

• Animales pequeños Dorsal nerve cord

Pharynx
with gill slits Stomach

Meiofauna o filtradores
Notochord
• (c) Heart

FIGURE 48.5
Tunicates (phylum Chordata, subphylum Urochordata).
(a) The sea peach, Halocynthia auranthium. (b) Diagram of the
structure of an adult tunicate. (c) Diagram of the structure of a
larval tunicate, showing the characteristic tadpolelike form. Larval
tunicates resemble the postulated common ancestor of the
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Kardong: Vertebrates: 2. Origin of Chordates Text © The McGraw−Hill 53

HISTORIA EVOLUTIVA DE LOS CORDADOS

Comparative Anatomy, Companies, 2009
Function, Evolution, Fifth
Edition

aligned, a pattern termed radial cleavage. At this point, the

TABLE 2.1 Fundamental Patterns in Bilateria
embryo is little more than a clump of dividing cells that soon
Development

Cordados provenientes de goes back to

• This effort to understand chordate origins Protostomes
Blastopore (mouth)
the beginning of the nineteenth century. One such view
artrópodos
traces chordate origins back to annelids and arthropods.
Spiral cleavage
Schizocoelic coelom
Gut
Ectodermal skeleton
Chordates from Annelids and Arthropods Nerve cord
Deuterostomes
Blastopore (anus)
Radial cleavage
Enterocoelic coelom
Mesodermal skeleton
become arranged into a round, hollow ball, with cells forming
the outer wall around a fluid-filled cavity within. One wall of
this ball of cells begins to indent and grow inward, a process
called gastrulation. The opening into this indentation is the
blastopore, and the indented cells themselves are destined to
become the gut of the adult. Indentation continues until cells
reach the opposite wall, where they usually break through,
forming a second opening into the primitive gut (the blasto-

Cordados
The first person toprovenientes debody
anélidos
pore being the first). The now multicellular embryo is com-
• propose that the chordate plan was
Embryonic development; details of early cleavage (p. 161)
posed of three basic tissue layers: an outer ectoderm, an inner
In both bilaterian groups, the egg begins to divide endoderm that forms the lining of the gut, and a mesoderm
Gut after fertilization, a process termed cleavage, until
derived from a flipped-over version of an arthropod’s was repeatedly
Nerve
the very young embryo cord
is made up of many cells formed from
that forms the layer between the two. If a solid mass of meso-
Annelid/Arthropod
dermal cells splits to form the body cavity within them, the
Geoffroy Saint-Hillaire, French zoologist. In 1822 he put the original single-celled egg (figure 2.3). In some animals, result is a schizocoelom (figure 2.3a). If, instead, the mesoderm
•forthArgumentación débil
this theory, perhaps inspired by his dissections of lob-
dividing cells of the embryo are offset from each other, a pat-
tern known as spiral cleavage. In others, the dividing cells are
arises as outpocketings of the gut that pinch off to form the
body cavity, the result is an enterocoelom (figure 2.3b).

sters, but also as part of his larger view that all animals
shared a common, underlying body plan that nature Coelom Mesoderm

Comparaciones sobre
Georgesanalogías
Protostome
•molded into dramatic variations. Cuvier led those Blastopore Mouth
Annelid
(earthworm)
Anus

opposing this theory, and in 1830 publicly debated Saint-

Hillaire. Cuvier ticked off a long list of differences that sur-
Segmentación
•passed the similarities, and y articulación
temporarily deissue
quieted the los (a) Spiral cleavage
Chordate Mouth develops
from blastopore
for most scientists. Coelom originating

apéndices vs segmentación muscular

Separate theories put forth later in the nineteenth
(a) Primitive
gut
from split of
mesoderm

century revived this view that annelids or arthropods might Arthropods

be chordate ancestors. In the early twentieth century, biolo- Chordate

•gist El
W. cordón nervioso
H. Gaskell and, shortly thereafter, William Patten
(modern bony fish)

Primitive Anus
revived the issue and presented a closely argued case in sup- Deuterostome
gut
Coelom

port of annelid and/or arthropod ancestry for chordates. Blastopore

• Desarrollo
Their reasoning,detaken
la boca
together, went as follows. Fish Echinoderm Mouth

Annelids and arthropods share with chordates similarities of (sea cucumber)

basic body design. All three groups are segmented. All

Formación
•exhibit delbrain
similarities in gross celoma y tipowith
regionalization, defore-
clivaje Amphibian
Radial cleavage
Coelom originating Anus
(b)
brain and hindbrain. Finally, the basic chordate body plan is
from outpouching
of gut

present in annelids and arthropods, although upside down Mammal Anus develops
from blastopore

(figure 2.29a,b). In annelids and arthropods, the nerve cord FIGURE 2.3 Protostomes and deuterostomes. Bilateria are divided into two major groups on the basis of embryonic

occupies a ventral position below the gut along with a major characteristics. (a) Protostomes usually show spiral cleavage, coelom formation by splitting of the mesoderm, and derivation of the mouth
(b)
from the blastopore. (b) Deuterostomes often exhibit radial cleavage, coelom formation by outpocketing of the gut, and derivation of the
 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Cordados provenientes de los equinodermos
kar24239_ch02_048-081.qxd 12/17/10 6:44 PM Page 76

• W. Garstand, afinidades embriológicas

FIGURE 2.30 Garstang’s theory
of the origin of the chordate body
• Hipótesis Auricularia plan. The proposed common ancestor of
chordates (left) was bilaterally symmetrical Circumoral Dorsal nerve
and had the external appearance of a ciliated band cord Pharyngeal
slits
young echinoderm larva. The ancestor’s
Mouth
circumoral ciliated bands and their
• Larva auricularia, Holoturios
associated underlying nerve tracts moved
dorsally to meet and fuse at the dorsal
Ciliated
adoral band
Endostyle

midline, forming a dorsal nerve cord in the Gut

chordate body plan. The adoral ciliated band

• Diplerula gave rise to the endostyle and ciliated tracts

within the pharynx of the chordate.
Scientists other than Garstang noted that Notochord
the appearance of pharyngeal slits improved
efficiency by providing a one-way flow for Anus
• Simetría bilateral the food-bearing stream of water. A
notochord appeared later, and with the
Dipleurula larva of echinoderm

swimming musculature, is a locomotor

advantage in the larger organism.

• Boca, banda adoral Chordate body plan

Garstang reasoned that because of these embryonic affini-
ties, echinoderms or a group very similar to echinoderms
• Banda circumoral, locomoción
were the likely chordate ancestors.
At first this seemed farfetched. Adult echinoderms, such
as starfish, sea urchins, sea cucumbers, and crinoids, offer little
to suggest a phylogenetic affinity with chordates. They have
tube feet, calcium carbonate plates in their skin, and pentara-
dial (five-armed) body symmetry. Let’s examine Garstang’s
they met at the midline together with the underlying nerve
tract, the antecedent of the nerve tube (figure 2.30).
Garstang pointed to the rolling up of the neural tube during
vertebrate embryology as an embryonic remnant of this
phylogenetic event. The segmental tail musculature with
notochord evolved simultaneously with the neural tube—
essentially a new nervous system serving to activate and
control the new muscularized tail. On the other hand,
 HISTORIA EVOLUTIVA DE LOS CORDADOS
kar24239_ch02_048-081.qxd 12/17/10 6:44 PM Page 76

• Cordados provenientes de los

equinodermos FIGURE 2.30 Garstang’s theory
of the origin of the chordate body
plan. The proposed common ancestor of

Similitudes bioquímicas en las proteínas chordates (left) was bilaterally symmetrical Circumoral Dorsal nerve
• and had the external appearance of a ciliated band cord Pharyngeal
slits
young echinoderm larva. The ancestor’s
sanguíneas circumoral ciliated bands and their
associated underlying nerve tracts moved
Mouth

• Química muscular: compuestos de fósforo
Ciliated Endostyle
dorsally to meet and fuse at the dorsal adoral band
midline, forming a dorsal nerve cord in the Gut
chordate body plan. The adoral ciliated band
gave rise to the endostyle and ciliated tracts
within the pharynx of the chordate.
Scientists other than Garstang noted that Notochord

Creatina en los vertebrados

the appearance of pharyngeal slits improved
• efficiency by providing a one-way flow for Anus
the food-bearing stream of water. A Dipleurula larva of echinoderm
notochord appeared later, and with the
swimming musculature, is a locomotor
• Arginina en los invertebrados advantage in the larger organism.
Chordate body plan

• Arginina y creatina en algunos

Garstang reasoned that because of these embryonic affini-
ties, echinoderms or a group very similar to echinoderms
they met at the midline together with the underlying nerve
tract, the antecedent of the nerve tube (figure 2.30).
were the likely chordate ancestors. Garstang pointed to the rolling up of the neural tube during
equinodermos al igual queAten tunicados
first this y echinoderms, such
seemed farfetched. Adult vertebrate embryology as an embryonic remnant of this
as starfish, sea urchins, sea cucumbers, and crinoids, offer little phylogenetic event. The segmental tail musculature with
hemicordados to suggest a phylogenetic affinity with chordates. They have notochord evolved simultaneously with the neural tube—
tube feet, calcium carbonate plates in their skin, and pentara- essentially a new nervous system serving to activate and
dial (five-armed) body symmetry. Let’s examine Garstang’s control the new muscularized tail. On the other hand,
hypothesis more closely. lengthening of the adoral band near the mouth and into
the pharynx provided the beginnings of an endostyle. The
Auricularian Hypothesis larva now lacked only pharyngeal slits to complete its trans-
Both echinoderms and chordates are deuterostomes that formation into a full-fledged chordate.
 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Hipótesis de Romeer,
Hemicordado colonial

• Estilo de vida de los pterobranquios

• Equinodermos y pterobranquios
relacionados con su ancestro

• Cordado ancestral no móvil, ni

bilateral

• Pruebas apuntan a un ancestro

sésil, béntico, suspensívoro

• Por eje: braquiopodos y briozoarios

 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Hipótesis de Romeer, Hemicordado colonial

• Cambio en la obtención del alimento, sustitución

del penacho

• Filtración por branquias

• Larva placntónica

• Balanogosus, larva ciliada

• Mejor larva parecida a un renacuajo, tunicados

• Posible un nuevo estilo de vida

• Conservaron las estructuras locomotoras

• Aparato filtrante faringeo

• Abandono del estado adulto sésil

 HISTORIA EVOLUTIVA DE LOS CORDADOS
GUERRERO Y GONZÁLEZ, (2.011)
• Aparición de registros fósiles, Cámbrico

• Inclusión del oxígeno

• Derrumbes, cambios en los ciclos biogeoquímicos de los suelos, mineralización

del agua

• Mineralización de los cuerpos de los organismos, incluidos los primeros cordados

• Dos faunas con fósiles

• Burges Shales (Columbia Británica, Canadá) Pikaia. 505 millones de años

• Chengjiang provincia de Yunnan, China. 520 millones de años

• Filum Vetulicolia primeros representantes de los Deuterostomados

 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Didazoon, Pomatrum, Vetulicola y
Xidazoon

• Cuerpo bipartito:

• Lado anterior voluminoso, lado

posterior segmentado (7
segmentos)

• Cinco bolsas branquiales, arcos

branquiales

• Mancha oscura posterior, endostilo

• Esqueleto interno
 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Swalla y Smith (2008)

• Ancestro de los cordados

• Organismo vermiforme segmentado

• Notocorda dorsal

• Nervios dorsales huecos

• Mucosa faringea, filtración

• Cola postanal

• Argumento no consensuado
 HISTORIA EVOLUTIVA DE LOS CORDADOS
• Yunnanozoons, cordado basal

• Semejante a un pez

• Cola postanal

• Notocorda

• Branquias

• Cefalización, con desarrollo de ojos

y cerebro limitado

• Miómeros laterales

• Posición taxonómica incierta

 HISTORIA EVOLUTIVA DE LOS CORDADOS

• Conodontos

• Alargados

• Cuerpo suave

• Paquetes musculares en V

• Ojos

• Denticulos conicos de
fosfato de calcio
HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS

PAOLA ESCOBAR RAMOS

BIÓLOGO, DOCENTE
MSC. CIENCIAS BIOLÓGICAS
 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS
• Registros fósiles escasos

• Vertebrados no comparables con los invertebrados actuales, especializados

• Divergencia, 500 millones de años

• El estado actual de los invertebrados, difícil de comparar

• Ancestros de los cordados extinguidos

• No ayudan los registros fósiles ni las pruebas genéticas

• Remodelaciones corporales

• Ventajas fisiológicas

• Adaptaciones favorables
 evolución de los vertebrados.
HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
Paedogenesis puede ocurrir en la evolución de los
vertebrados
• Paedogenesis Adultos conserva características
Paedogenesis: Adultos conserva características
que se observan en el desarrollo embrionario
que se observan en el desarrollo embrionario

Vertebrate

Adult Urochordate Ancient Cephalochordate

No Paedogenesis Paedogenesis

Ancient Urochordate larva

 SUBFILUM VERTEBRATA
48.1 Attaching muscles to an internal framework greatly improves movement.
• Animales complejos
he Chordates Pharyngeal Hollow dorsal
pouches nerve cord
hordates (phylum Chordata) are
uterostome
Ambientes acuáticos, terrestres
• coelomates whose near-
relations in the animal kingdom

Arthropods
Sponges

Flatworms

Annelids

Chordates
Nematodes
Cnidarians

Mollusks

Echinoderms
y aereos
the echinoderms, the only other
uterostomes. However, unlike
hinoderms, chordates are character-
Diferentes estilos de vida
d by a• notochord, jointed appendages,
d segmentation. There are some
000 species of chordates, a phylum
t includes birds, reptiles, amphib-
Características de los cordados
• and mammals.
s, fishes,
Four features characterize the chor-
es and have played an important
Al igual que los tunicados y
• evolution of the phylum
e in the
gure 48.2):
anfioxus Postanal
tail
1. A single, hollow nerve cord
Notochord
runs just beneath the dorsal sur-
face of the animal. In verte-
• Cinco características del filum
brates, the dorsal nerve cord differentiates into the
FIGURE 48.2
Some of the principal features of the
brain and spinal cord. chordates, as shown in a generalized
2. A flexible rod, the notochord, forms on the dorsal embryo.
side of the primitive gut in the early embryo and is
 SUBFILUM VERTEBRATA: COLUMNA Y
CRÁNEO
48.3 The vertebrates have an interior framework of bone.
• La notocorda es reemplazada
Characteristics
durante el desarrollo of Vertebrates
Ectoderm
Vertebrates (subphylum Vertebrata) are chordates with a
embrionario
spinal column. The name vertebrate comes from the indi- Neural tube
vidual bony segments called vertebrae that make up the
Notochord
spine. Vertebrates differ from the tunicates and lancelets in
• Columna vertebral ósea
two important respects:
1. Vertebral column. In vertebrates, the notochord is
replaced during the course of embryonic develop-
• Segmentos que cubren y
ment by a bony vertebral column. The column is a
series of bones that encloses and protects the dorsal
Vertebral
body developing
Forming
protejen el nervio dorsal
nerve cord like a sleeve (figure 48.8).
2. Head. In all vertebrates but the earliest fishes, there
around notochord
neural arch
Blood vessels
is a distinct and well-differentiated head, with a skull
and brain. For this reason, the vertebrates are some-
• Cabeza diferenciada: esqueleto
times called the craniate chordates (Greek kranion,
“skull”).
y Incerebro
addition to these two key characteristics, vertebrates
differ from other chordates in other important respects: Rib Neural arch

1. Neural crest. A unique group of embryonic cells

• Cordados craneados
called the neural crest contributes to the development
Centrum

of many vertebrate structures. These cells develop on

the crest of the neural tube as it forms by an invagina-
tion and pinching together of the neural plate (see
 SUBFILUM VERTEBRATA: COLUMNA Y
CRÁNEO
48.3 The vertebrates have an interior framework of bone.
• Crestas neurales,
Characteristics of Vertebrates
Ectoderm
Vertebrates (subphylum Vertebrata) are chordates with a
• Células embrionarias
spinal column. The name vertebrate comes from the indi-
vidual bony segments called vertebrae that make up the
Neural tube

Notochord
spine. Vertebrates differ from the tunicates and lancelets in
two important respects:
• Derivan del tubo
1. Vertebral column. neural
In vertebrates, the notochord is
replaced during the course of embryonic develop-
ment by a bony vertebral column. The column is a Vertebral

• Desarrollan otras estructuras

series of bones that encloses and protects the dorsal
nerve cord like a sleeve (figure 48.8).
body developing
around notochord
Forming
neural arch
2. Head. In all vertebrates but the earliest fishes, there
en los vertebrados
is a distinct and well-differentiated head, with a skull
Blood vessels

and brain. For this reason, the vertebrates are some-

times called the craniate chordates (Greek kranion,
“skull”).
• ENDOESQUELETO, fosfato
In addition to these two key characteristics, vertebrates
de calcio
differ from other chordates in other important respects: Rib Neural arch

1. Neural crest. A unique group of embryonic cells Centrum

called the neural crest contributes to the development
of many vertebrate structures. These cells develop on
the crest of the neural tube as it forms by an invagina-
tion and pinching together of the neural plate (see
 SUBFILUM VERTEBRATA: COLUMNA

• Vertebrados, nombre del grupo

• Vertebras, serie de bloques de huesos y cartílagos

unidos

• Espina dorsal

• Lo anterior define el eje corporal

• Discos intervertebrales, entre las vértebras

 kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 84

SUBFILUM VERTEBRATA: COLUMNA

VERTEBRAS

• Cuerpo cilindrico o centro

• Notocordo

• Arco neural

• Médula espinal
Neural spine Dorsal hollow
nerve cord
• Arco hemal Intervertebral Neural arch
body
Centrum
• Vasos sanguíneos
Hemal arch Blood vessel
• Primeros vertebrados, sin vertebras
Hemal spine
pero con notocordo fuerte

FIGURE 3.2 Basic vertebra. Vertebrae replace the

 SUBFILUM VERTEBRATA: CRÁNEO

• La mayor innovación, cranio: craniata o craneados

• Compuesta por hueso y cartilago

• Órganos sensoriales, protege al cerebro y cerebelo

• Tubo neural se alarga (bulbos)

• Encéfalo anterior definido

 Ectoderm

Mesoderm

(a)

SUBFILUM VERTEBRATA: CRÁNEO Endoderm Gastrocoel

• Placas epidermales y Neural fold

Epimere
crestas neurales, caracteres Mesomere

diagnósticos del grupo en Mycoel

Somatic
mesoderm
estados embrionarios
Nephrocoel
Hypomere
Splanchnic
4239_ch05_161-211.qxd 12/21/10 7:36 PM Page 170 (b) mesoderm
Coelom

• No observados en los
ancestros
Neural crest Neurocoel

Neural tube
Chordamesoderm
Organos sensoriales y
Dermatome
• Somite Myotome
Neural plate
algunos dientes
Dorsal mesentery
Blastocoel Sclerotome
Early
GastrocoelSomatopleure
(c) Gut tube cavity
P Splanchnopleure

• Células embrionarias P Ventral mesentery

FIGURE 5.6 General steps in successive

(a)
P differentiation of the mesoderm and neural tube.
Gastrocoel
Mesoderm initially comes to lie between the other two germ
layers (a), and differentiates into three major regions: the epimere,
Blastopore the mesomere, and the hypomere (b). Each of these gives rise to
SUBFILUM VERTEBRATA: Característica diagnósticas
de los cordados en alguna etapa de su vida

• Tegumento con modificaciones

• Epidermis

• Surge del ectodermo

• Epitelio estratificado

• Dermis

• Surge del mesodermo

• Tejido conectivo
SUBFILUM VERTEBRATA: Característica diagnósticas
de los cordados en alguna etapa de su vida

• Faringe musculosa

• Sacos faringeos, abren hacia afuera como hendiduras

y contienen agallas

• Sacos faringeos son fuentes de otras glándulas

• Esqueleto axial: cráneo, columna, esternón y costillas

• Esqueleto apendicular: apêndices delantero y trasero,

cinturon pectoral y pélvico
 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: LA CONQUISTA DEL AGUA
La Conquista del Agua
HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
LA CONQUISTA DEL AGUA

• Prevertebrados, parecidos al anfioxo

• Cilios

• Agnathos

• Sin mandíbula con bomba muscular

• Gnathostomados

• Con mandíbula
HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 85

LA CONQUISTA DEL AGUA, Prevertebrados

• Surgieron de los protochordados

• Dos puntos de vista contrastante Gnathostome

Adult
Larva
• Depredadores incipientes
Jaws
• Filtradores Agnathan
(grasping)

Egg
• Comportamiento filtrador Adult

(maquinaria ciliar) comun de Prevertebrate

Vertebrate

hemichordados, Muscular pump

(suspension feeding)
cephalochordados y
urochordados
Cilia-mucus
(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 85

LA CONQUISTA DEL AGUA, Prevertebrados

• Similar al Amphioxus

• Nadadores libres Gnathostome

Adult
• Tolerantes a ambientes de Larva

estuarios
Jaws
Agnathan
(grasping)
• Cambio a vertebrados: cambios Egg
mecánicos Adult

Vertebrate
Prevertebrate
• Faringe: bomba muscular Muscular pump
(suspension feeding)

• Bandas circulares musculosas

• Colágeno por cartílago Cilia-mucus

(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 85

LA CONQUISTA DEL AGUA, Agnathos

• Contraccion de la faringe,
expulsión del agua
Gnathostome

• Cartilagos, la faringe Adult

Larva

recuperaraba su forma inicial

Jaws
Agnathan
(grasping)
• Bomba ciliar reemplaza a la Egg

maquinaria ciliar Adult

Vertebrate
Prevertebrate
Muscular pump
• Maquinaria ciliar insuficiente (suspension feeding)

en la captación del alimento

para los animales mas
grandes Cilia-mucus
(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
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LA CONQUISTA DEL AGUA, Agnathos

• Contraccion de la faringe,
expulsión del agua
Gnathostome

• Cartilagos, la faringe Adult

Larva

recuperaraba su forma inicial

Jaws
Agnathan
(grasping)
• Bomba ciliar reemplaza a la Egg

maquinaria ciliar Adult

Vertebrate
Prevertebrate
Muscular pump
• Maquinaria ciliar insuficiente (suspension feeding)

en la captación del alimento

para los animales mas
grandes Cilia-mucus
(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
kar24239_ch03_082-127.qxd 12/21/10 7:27 PM Page 85

LA CONQUISTA DEL AGUA, Agnathos

• Branquias

• Órganos respiratorios Gnathostome

complejos Adult
Larva

• Tapizados por vasos Jaws

sanguíneos Agnathan
(grasping)

Egg

• Agua + alimento + oxígeno Adult

Vertebrate
Prevertebrate
Muscular pump
• Incrementa la resistencia (suspension feeding)

• Nueva faringe: paso del agua a

través de las nuevas branquias Cilia-mucus
(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

HISTORIA EVOLUTIVA DE LOS VERTEBRADOS:
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LA CONQUISTA DEL AGUA, Gnagthostomados

• Cambio en el método de
alimentación
Gnathostome
• Captura fortuita Adult
Larva

• Expanción de la bomba faringea

Jaws
Agnathan
(grasping)
• Cierre de la boca Egg
Adult
• Primeros vertebrados Prevertebrate
Vertebrate

succionadores (ineficiente) Muscular pump

(suspension feeding)

• Aparición de mandíbulas:
expanción faringea y succión mas
fuerte Cilia-mucus
(suspension feeding)

FIGURE 3.3 Origin of vertebrates. A more active,

 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA
• Del agua a la tierra, importante para los animales

• Invasión del habitat, especies peligrosas

• Vida desarrollada en el agua

• Composición de los animales

• Actividades celulares y fisiología

• Transición a la tierra, requiere de modificaciones

• Corporal

• Contenido de oxígeno

• Densidad

• Regulación de la temperatura

• Diversificación del habitat

 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA

• Corporal

Clack 2009, The Fish–Tetrapod Transition:

New Fossils and Interpretations.
 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA

• Corporal
 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA
• Contenido de oxígeno

• 20 veces mas abundante en el aire

• Difunde con mayor rapidez

• Obtención fácil, pulmones

• Densidad

• 1000 veces menos densos

• 50 veces menos viscosos

• Soporte contra la gravedad

• Extremedidades y remodelación del esqueleto

 HISTORIA EVOLUTIVA DE LOS
VERTEBRADOS: DEL AGUA A LA TIERRA
• Regulación de la temperatura

• Ambiente terrestre

• Ciclos de temperatura, congelamiento, derrite, sequias, lluvias

• Comportamientos y estrategias fisiológicas

• Homeotermia, regulación constante de la temperatura corporal

• Animales ectotérmicos o poiquilotérmicos y endotermia

• Diversificación del habitat

• Polos, coníferas, bosques tropicales, sabanas, desiertos, islas oceánicas y

montañas

• Protección para un eficaz desarrollo