REPTILES

PAOLA ESCOBAR RAMOS

MSC. CIENCIAS BIOLÓGICAS
BIÓLOGA
2.016
 Sauropsida Synapsida
ing time. But in animals that cannot walk immediately, the amphibia
pelvis is not fully hardened at birth. Of the five dinosaur Tihen be
Reptilia species for which embryos have been found, all had bony predation
that in th
Parareptilia Eureptilia have been

AMNIOTAS
the speci
Diapsida FIGURE 7.4
branes) h
even mod
Egg
Archosauromorpha Lepidosauromorpha
gilled la
Developing brain
Amnion Embryo Allantois
cosauriscu
Archosaurs of gilled
ians even
"Euryapsids"
logenetic
• Embriones con membrana Dinosauria Wer

Crocodylomorpha
Diadectomorpha

to deposi
extraembrionarias y cáscara

Sauropterygia
"Thecodonts"
primarily

Ichthyosauria

Lepidosauria
Captorhinids
Stem groups

Stem groups

Ornithischia
response
Mesosaurs

Pterosaurs
Testudines

Saurischia
calcárea to protec
These qu

Aves
Synapsida
Ancient
• Amniotas radiaron de dos Chorion Yolk sac Shell Albumin
Air
space
Reptiles
most of
líneas evolutivas: ?
Generalized structure of the amniotic egg. Its membranes—chorion,
amnion, yolk sac, and allantois—protect the embryo and provide it
and aeria
as did ca
with metabolic support. rise to th

• Saurópsidos y Sinapsidos Sauropsida Synapsida

Parareptilia Eureptilia
• Saurópsidos: reptiles y aves

• Sinápsidos: mamíferos

(a) Phylogeny (b) Major trends

FIGURE 3.27 Amniotes, phylogenetic relationships. (a) Phylogeny of major groups. Note the major trends within amniotes,
 AMNIOTAS
562 PART 3 The Diversity of Animal Life

Amniota
Sauropsida
Synapsida Anapsida Diapsida
Lepidosauria Archosauria
Squamata

Lizards and
Mammals Turtles Sphenodon Amphisbaenids snakes Crocodilians Birds

Squamata: fusion of snout Archosauria: presence of

Orbit bones, characteristics of opening anterior to eye,
palate, skull roof, vertebrae, orbit shaped like inverted
ribs, pectoral girdle, humerus triangle, teeth laterally
compressed
Lepidosaurians: character-
istics of bone growth,
skull, pelvis, feet
Dorsal
temporal
Anapsid skull Testudines: solid-roofed anapsid opening Orbit
skull, plastron, and carapace
Lateral derived from dermal bone and
temporal fused to part of axial skeleton Lateral
opening Orbit temporal
Diapsids: diapsid skull opening
Synapsids: skull with single with 2 pairs of temporal
pair of lateral temporal openings
openings Diapsid skull
Turtle-diapsid clade (Sauropsida) Figure 28-2
characteristics of skull
Cladogram of the living Amniota showing monophyletic groups. Some of the
and appendages
Synapsid skull shared derived characters (synapomorphies) that are diagnostic for the
lineages are given. The skulls represent the ancestral condition of the three
Amniotes: extraembryonic groups. The skulls of modern diapsids and synapsids are often highly modified
membranes of amnion,
by loss or fusion of skull bones that obscures the ancestral condition.
chorion, and allantois
Representative skulls for anapsids are Nyctiphruetus of the upper Permian; for
diapsids, Youngina of the upper Permian; for synapsids, Aerosaurus, a

• Relaciones entre los grupos de amniotas: pelycosaur of the lower Permian. The relationships expressed in this cladogram
are tentative and controversial, especially that between birds and mammals.
Contrary to the view shown here, in which mammals are the outgroup, some
authorities support a sister-group relationship between birds and mammals
based on several kinds of molecular and physiological evidences.

• Fenestraciones craneales
 REPTILES
kar24239_ch03_082-127.qxd 12/21/10 7:28 PM Page 113

absent. Snakes lack both upper and lower temporal bars.

Because these connecting constraints are deleted in the
skulls of lizards and snakes, both of these vertebrate
groups, especially snakes, have increased jaw mobility that
kar24239_ch03_082-127.qxd 12/21/10 7:28 PM Page 112 enhances their ability to capture and swallow prey.

(a) Ichthyosaur Functional consequences of the loss

of temporal arches (p. 268)

Pararreptiles y Eurreptiles
The squamates include snakes, lizards, and a group
• of tropical or subtropical reptiles, the amphisbaenids.
Some taxonomists place amphisbaenids with lizards; oth-
specialized turtle body plan. The most primitive turtles for ers treat them as a distinct group. All amphisbaenids are
burrowers; most are limbless and prey upon arthropods
which there is a fossil record (b) (Late
Plesiosaur Triassic) already pos-
(figure 3.33b). The majority of living squamates are lizards
sessed this shell housing limbs within. Where fossils fail to
• Pararreptiles: Testudinos clarify, modern molecular
kar24239_ch03_082-127.qxd

mechanism—the fundamental
FIGURE
genetics
12/21/10
(a) Ichthyosaur,
3.327:28
asuggests
Marine
porpoise-like
changea plesiosaur,
(b) Sauropterygian,
PM

in a fewabout
reptiles
thereptile
Page

turtle
of the Mesozoic.
underlying
111
about 1 m long.
7 mHox
in length.
or snakes (figure 3.33c,d). Many people are surprised to
learn that some species of lizards (other than amphis-
baenids) are limbless, like snakes; therefore, the presence

(tortugas) cráneo anapsido genes—is the apparent(a)basis for the radical

After Romer; (b) after Andrews. transformation Eocaptorhinus or absence of limbs alone does not distinguish 1 cm snakes from
of the amniote body plan into the unique design of turtles. true lizards. Instead, differences in internal skeletal
FIGURE 3.31 anatomy,Captorhinomorph.
especially in the skull, Skeleton are used to of diagnose
the the
Turtle axial
teethskeleton
around the (p. rim
315)of a beaklike mouth testify reptile
to an two groups.
Eocaptorhinus, fromFurther,
the Permian.lizardsAhave North movable
American eyelids and most
active predaceous lifestyle. Preserved in fine-grained member sedi- of thehave an external
captorhinid family.auditory meatus (opening). Snakes lack
Modern turtles belong ments, either
ichthyosaur to the Pleurodira
stomach contents or include
to prodigious Reptilia both structures.

Eurreptiles: Squamata to fish,Afterand Heaton and Reisz.

the Cryptodira, depending on the method (squidlike
they employ
• retract their head into their
amounts
some,shell.
of belemites
Pleurodires
hatchling
mollusks),
flex their neck
turtles. Relatively huge eyes gave them
in
The taxon Reptilia, in a restricted sense, applies to the
Archosauromorpha Encompassed within the archosauro-
Parareptilia and Eureptilia, which together share similarities of
light-gathering power
laterally to retract the head, whereas cryptodires flex their for dim waters and also the ability to morphs are several stem groups, small assemblages of diapsids
the braincase
knownthat fromdistinguish them from
largethe mesosaurs. The
(lagartos y serpiente) fine-focus on small,
neck vertically. These two groups seem to share a common quick prey. Fossil
female show fully-formed young ready for birth or in the
ancestor, Proganochelys,process, from evidence
the lateforTriassic. The term
remains of a pregnant
eureptilians
digit and
kar24239_ch03_082-127.qxd
live birth (not egg laying) in these
taxon
ankle
characteristic
Anapsida
for 12/21/10
the and
was
articulations,
second.birds.Anapsida
7:28
fossils,
PM Page 114
once long
Archosaurs
andand
used
and
a very
for slender
this first
similarities
weredisplay
reptiles
group,
group
which includes familiar forms such as crocodiles, dinosaurs,
limbs,
diagnosed
a trend
thesimilar
and
of toward
archosaurs,
Diapsida
bonebystruc- skulls
increasing
tortoise is sometimes applied marinetoreptiles
turtles restricted
(see figure to1.33). land,One but of theture within
without the skull. Captorhinid
temporal fenestrae; reptiles
Diapsida by were
skullsThe small, about
withforelimbs
two tem-tend
Crocodylia (caimanes y
largest bipedalism, or two-footed locomotion.
no formal taxonomic
FIGUREdistinction
3.29
ichthyosaurs is made
rivaledbetween
Mesosaur. This aquatic
or exceeded turtle theand
amniote livedofina 20
size cm poral
moderm in length,to beand
fenestrae. generally
However,
reduced, whereas similar
analysis oftomulticharacters,
the hindlimbs modern are drawnlizards com-
under in the
tortoise. the middle Permian.The
sperm whale.long tail was used for swimming, and the pared
that they hadbody to just skull
a well-ossified temporal
to become the region,
skeleton reveals a
(figure 3.31).and
major weight-bearing slightly different
Rows of
locomotor
limbs were probably paddle shaped. Overall length, about 1 meter. evolutionary history.The
appendages. Some skullreptiles with
is diapsid, anapsid
but skullsacross
an additional dofenestra
not
babilla) Aves. Cráneo
Traditionally, turtles are viewed as the only living rep- tiny, sharp teeth along the margins of the jaws and
Sauropterygia Sauropterygians were, along with opens Anapsida;
on the facecaptorhinids
resentatives After McGregor; von
of the earliest Huene.
reptiles, a sister group to diapsids, the roof occur within
of the mouth as wellbetween as an agile maxilla
have and
bodyanapsidlacrimal
suggest skulls,bones,
that but the
ichthyosaurs, the other Mesozoic lineage of diapsids spe- do fall antorbital
within the fenestra,
old as wellToday,
“diapsids.” as a mandibular
anapsida fenestratoina the
applies
as I treat them here (figure 3.27).
cialized to However,
an aquatic Imode should concede
of life. Their current insects
place- mightlower havejaw. been a major part of their diet, as they

diapsido
skull type, but it is used less as a taxon name. Taxonomically,
that some recent morphologicalment, evidence,
tentatively, that includes
is within fos-
the lepitosauromorpha. are in the similarly Thedesigned small,meaning
modern lizards. Cap-
to include all amniotes and their sister group, the diadecto- This Diapsida is now terminarchosaur,
used a more restricted “ruling
sense for reptile,”
a mono- recog-
sils, favors placement of turtles
group
morphs (figure 3.27).
within
includes the
the diapsid
early radiation,
nothosaurs (Triassic)torhinids
and
(a) the arenizes
Sphenodon broadly similar to anthracosaurs,
the extraordinary radiation and preeminence
phyletic lineage within the Eureptilia (figure 3.27). Note that
but cap- of this
later plesiosaurs
not primitively within reptiles generally. (Jurassic-Cretaceous)
Although currently that evolved from possess
torhinids group reptilian
during the features, Mesozoic. such
groupingas
Formally strong
as well. Thisjaw
then, archosaurs
birds get scooped up in this (c) cladistic sim-
unresolved, the outcome them.
of The
this plesiosaur
debate is body
important was heavy,
because the neck often
musculature, include
and “thecodonts,”
reptilian the
structural most
Sceloporus primitive
details in of the group,
their skull, croc-
Sauropsids long, and the limbs were modified into paddles that acted ply recognizes that birds are a natural but specialized
odiles, birds, pterosaurs, and two large groups, the Saurischia derivative
if turtles turn out to be diapsids, then a major revision would limbs, of and vertebral
earlier reptiles. column.
We return The firstlater,
to birds captorhinid occupied
Mesosaurs like oars or hydrofoils to propel the animal in water and Ornithischia. Taken together, but first we
Saurischia complete
and Ornithis-
be in order in comparative studies.
(figure 3.32b). First, turtles (if diapsids) tree stumps
our away
review of from standing
sauropsids. bodies of
chia constitute what laypeople informally think of as the water, providing
could no longer be assumed
The mesosaurs were the tofirstrepresent the ancestral
of many sauropsids to embark onadditional testimony “dinosaurs.”that they exploited
However, the land They
birds are descendants. further, evolved
a specialized
reptilian condition against aquatic
Lepidosauria existence
which other Modern (figure
derivedsnakes,3.29).
groups Fossils
could
lizards, are few,
Sphenodon, more and their Parareptilia
characteristic
within the Testudines
ofdinosaur (turtles)
reptilesradiation and
than of(figure
(d) Contia
an assortment
their3.27) earlyso,tetrapod of fos-they
formally,
be compared. and this puzzling
Second, ancestors
the group
absence exhibits
constitute the
of temporal nolepidosaurs.
close affinity
fenestrae into Permian/Early
A late otherancestors. sil groups (e.g.,
should be Pareiasaurus)
included. are included in the Pararep-
aquatic Triassic
sauropsids. group
The of
earliestlepidosaurs,
turtles (again, if diapsids) would imply a secondary condi- date to the
the late Permian,
Eosuchia, are most
but likely
tilia. They haveThe a distinctive
thecodonts ear
aroseregion
late in wherein
the Permian the eardrum and pros-
a much theappearance
earlier ancestors of is all modernbecause
suspected lepidosaurs.
of the Sphenodon,
primi- the
is
(b) Amphisbaena peredby
supported during
the the Triassic.(rather
squamosal Before becoming
than by extinct
the by the end
quadrate)
tion in which the two fenestrae
tuatara,
close up again, not an ances- Eureptilian Radiation Within the eureptilian radiation,The-
tive skull that lackedistemporal
the sole survivor
fenestrae. of The
a once widespread
elongate snout
FIGUREMesozoic
3.33and by of
thethe
Lepidosaurs. Triassic,
(a)retroarticular
Sphenodon. theyprocess,
gave
(b) Amphisbaenian, risea lepidosaur.
a burrowing to all later
backward archosaurs.
projection
(c) Lizard of (Contia).
(Sceloporus). (d) Snake
tral condition. Third, this would leave the Parareptilia there were three major lineages, all built on a diapsid design
 FIGURE 6.12 Amphibian skin. (a) Section through an
adult frog skin.A basal stratum basale and a thin, superficial
stratum corneum are present.The transitional layer between
them includes a stratum spinosum and a stratum granulosum.

TEGUMENTO
(b) Diagrammatic view of amphibian skin showing mucous and
poison glands that empty their secretions through short ducts to
the surface of the epidermis.

• Refleja la existencia terrestre

• Queratinización extendida

• Menos glándulas Hinge Hinge

Epidermis
Dermis

• Repliegue de la epidermis
(a)

• Union entre escamas, FIGURE 6.13 Reptile skin. (a) Epidermal scales. Exten
even along the body of the same individual. Snake body scales (to
bisagra flexible scales is a thinned area of epidermis, a “hinge” allowing skin flexi
shed, the basal cells produce an inner epidermal generation.Whi
• Escama grande en forma de from old outer epidermis.
(a) After Maderson; (b) after Landmann.
lámina, escudo
220 Chapter Six
 TEGUMENTO
hic09617_ch28.qxd 6/7/00 1:10 PM Page 564

564 PART 3 The Diversity of Animal Life

• Escama, cresta o espina

• Con hueso dérmico Scale

Epidermis
• Placas de hueso dérmico
Dermis
• Osteodermos, variables
entre los grupos
Osteoderm Melanophores Flexible hinge
• Bajo las escamas Figure 28-3
Section of the skin of a reptile showing the overlapping epidermal scales.
epidérmicas
Embryo
 serving cutaneous respiration.
hic09617_ch28.qxd 6/7/00 1:10 PM Page 564
Stratum Leydig cell
corneum Reptiles
Transitional The skin of reptiles reflects their greater commitment to a

TEGUMENTO
layer Epidermis
terrestrial existence. Keratinization is much more extensive,
Stratum and skin glands are fewer than in amphibians. Scales are
basale 564
Chromatophore PART 3 The Diversity of Animal Life
present, but these are fundamentally different from the der-
Mucous
Poison
mal scales of fishes, which are built around bone of dermal
gland Dermis
gland origin. The reptilian scale usually lacks the bony undersup-
port or any significant structural contribution from the der-
mis. Instead, it is a fold in the surface epidermis, hence, an
epidermal scale. The junction between adjacent epidermal
• Epidermis con estratos basal,
(b) Scale
scales is the flexible hinge (figure 6.13a). If the epidermal
scale is large and platelike, it is sometimes termed Epidermis
a scute.
granuloso y córneo
FIGURE 6.12 Amphibian skin. (a) Section through an
adult frog skin.A basal stratum basale and a thin, superficial
Additionally, epidermal scales may be modified into crests,
stratum corneum are present.The transitional layer between spines, or hornlike processes.
them includes a stratum spinosum and a stratum granulosum. Although not usually associated with scales,Dermis
dermal
• Dermis: tejido conectivo
(b) Diagrammatic view of amphibian skin showing mucous and
poison glands that empty their secretions through short ducts to
bone is present in many reptiles. The gastralia, a collection
of bones in the abdominal area, are examples. Where dermal
fibroso
the surface of the epidermis. bones support the epidermis, they are called osteoderms,
Osteoderm Melanophores Flexible hinge
Figure 28-3
• Muda o ecdisis, Section of the skin of a reptile showing the overlapping epidermal scales.

desprendimiento de piel Embryo

cornificada Layers:
Leathery shell
Outer
Hinge Inner
• Tortugas y cocodrilosHinge Epidermis
Chorion
Dermis Basal

modesta
(a)
Allantois
Rest Renewel Shed Rest
(b)

FIGURE 6.13 Reptile skin. (a) Epidermal scales. Extent of projection and overlap of epidermal scales varies among reptiles and
• Serpientes y lagartos, Yolk sac
even along the body of the same individual. Snake body scales (top) and tubercular scales of many lizards (bottom) are illustrated. Between
scales is a thinned area of epidermis, a “hinge” allowing skin flexibility. (b) Skin shedding. Just before the old outer layer of epidermis is
abandono de la piel
shed, the basal cells produce an inner epidermal generation.White blood cells collect in the splitting zone to promote separation of new
from old outer epidermis.
 TEGUMENTO
• Glándulas:

• Femorales: lagartos

• Odoríficas: tortugas y cocodrilos

• Odoríficas en la cloaca y glándulas en la

mandíbula inferior: caimanes

• Función reproductora
 REPRODUCCIÓN
• Ovíparos u ovoviviparous

• Construcción de nidos

• Nidadas dependientes de la
temperatura

• < 30Cº hembras

• > 34 Cº machos

• >26Cº < 36Cº muerte

embrionaria
 Mammals Turtles

REPTILES: Testudinos
Orbit

• Triásico

• Caparazón dorsal Anapsid skull

Lateral
• Costilla extendidas temporal
opening Orbit

Synapsids: skull
pair of lateral t
• Placas dérmicas opening

superficiales (escudos)
Synapsid skull

• Plastrón ventral

• Pieza córnea fusionada

 Mammals Turtles

REPTILES: Testudinos
Orbit

• Caparazón: dos capas

• Externa, capas de keratina. Anapsid skull

Lateral
• Capas nuevas crecen debajo temporal
opening Orbit

de las viejas Synapsids: skull

pair of lateral t
opening

• Interna, hueso
Synapsid skull

• Fusión de costillas, vértebras

y elementos dérmicos
osificados
 Mammals Turtles

REPTILES: Testudinos
Orbit

• Caparazón

• Encierra la cintura escapular y

Anapsid skull
pélvica
Lateral
temporal
• Reduce la vulnerabilidad opening Orbit

Synapsids: skull
pair of lateral t
• Previene heridas incidentales opening

• Reducen la mortalidad Synapsid skull

• Longevidad

• Almacén de alimentos

• Mayor masa corporal

 Por otra parte en los pleurodiros se ha fusionado la cintura pélvica a la
concha en tanto que en los criptodiros, la cintura pélvica se une a esta

REPTILES: Testudinos
por medio de ligamentos. Los pleurodiros abarcan tres familias vivientes:
Pelomedusidae, Podocnemididae y Chelidae, en tanto que la mayor parte
de las 313 especies de tortugas vivientes, 77 géneros y 11 familias son
criptodiros (>200 especies). De las 14 familias de Testudines, hay repre-
sentantes de 9 en la región tropical andina cubierta por este libro (ERNST
& BARBOUR 1989; FRITZ & HAVAŠ, 2006).

Por otra parte en los pleurodiros se ha fusionado la cintura pélvic

concha en tanto que en los criptodiros, la cintura pélvica se une
por medio de ligamentos. Los pleurodiros abarcan tres familias vivi
• Tortugas actuales: Pelomedusidae, Podocnemididae y Chelidae, en tanto que la mayor
de las 313 especies de tortugas vivientes, 77 géneros y 11 familia

Pleurodiros y Criptodiros criptodiros (>200 especies). De las 14 familias de Testudines, hay r

sentantes de 9 en la región tropical andina cubierta por este libro (E
Vista posterior de dos conchas que muestran: Arriba a la cintura pélvica
& BARBOUR 1989; FRITZ & HAVAŠ, 2006).
(flexión del cuello)
fusionada a la concha (Pleurodiros) y abajo a la cintura pélvica unida con
ligamentos (Criptodiros) fotos A. González.

La concha de las tortugas (la cual incluye el espaldar o carapax en la parte

dorsal y la porción aplanada del vientre denominado plastrón o peto), se
encuentran unidas, a los lados y entre sí por un área llamada puente. La
• Vertical, Criptodiros concha consta de dos elementos: un escamado epidérmico córneo (escudos)
que reviste una armadura ósea de origen dérmico, dispuestos a manera de
un mosaico característico. La gran mayoría de las tortugas vivientes poseen
54 escudos epidérmicos de los cuales 38 recubren el carapax y 16 están
ubicados sobre el plastrón y si bien no todas las especies de tortugas poseen
• Lateral, Pleurodiros el mismo número de escudos, este patrón es esencialmente el mismo en

80 • Tortugas y Crocodylia

Vista posterior de dos conchas que muestran: Arriba a la cintura pélvica

fusionada a la concha (Pleurodiros) y abajo a la cintura pélvica unida con
ligamentos (Criptodiros) fotos A. González.

La concha de las tortugas (la cual incluye el espaldar o carapax en la

dorsal y la porción aplanada del vientre denominado plastrón o pet
 nsible, a su vez no juega se aprecia una relación inversa. Generalmente, en la mayoría de tortugas
de sustancias distantes. Claveel para
acuáticas, las hembras tienen tamaño lascorporal
especies más de tortugas
grande. La cópula
ar tienen una membrana se lleva a cabo con el macho montado sobre el carapax de la hembra, por
dulceacuícolas de la familia Chelydridae
REPTILES: Testudinos
ue al vibrar transmite los

rían entre las diferentes

lo que el plastrón de estos es cóncavo, especialmente en aquellas especies
que tienen caparazones muy
que distingue los sexos es que
arqueados.
Tamaño grande, longitud
los machos
el caparazón;
Ladelcaracterística
poseenmuyuna
plastrón cruciforme
más
caparazón hasta 45 cm,
colacabeza
reducido;
mandíbulas...............Chelydra acutirostris
colaconstante
casi tan larga como
másgrande
larga,
y concon
poderosas

de agua disponible en el la apertura cloacal ubicada en una posición más distal. En las tortugas
das las especies acuáticas marinas los machos tienen las uñas más desarrolladas sobre las aletas
os productos nitrogena-
), la cual requiere el paso igual forma en muchas tortugas dulceacuícolas las uñas de los dígitos del
menos tóxica y soluble en pie de los machos son más grandes y curvas para aferrarse a los lados
del caparazón de las hembras; tal vez con el mismo propósito funcionen
Tortuga terrestre
uctos varia.
• Las tortugas
o y altos porcentajes de
urea y las que habitan en muchos kinostérnidos. Aún cuando la mayor parte de las tortugas son
tugas de las Caparazón alto y curvo
• zonas áridas mudas, algunas especies son capaces de producir sonidos bastante notables
antidad de agua posible, durante el apareamiento como el caso de Chelonoidis spp.
, el cual es insoluble en
Plastrón amplio
lo regular •estas tortugas
nes de agua, casi limpia.
e sus cuerpos a través de
•
e los ojos.Tortuga acuática
de glándulas de almizcle
del puente,•muy cerca de
Caparazón plano
ucturas producen secre-
as interacciones sociales
dadores. Otras Plastrón reducido
• tortugas
enen glándulas sobre el Vista ventral y dorsal de un adulto de Chelydra acutirostris - Zoológico de Cali,
Colombia, fotos C.A. Galvis
porada reproductiva. Cópula de la tortuga terrestre morrocoy - Chelonoidis carbonaria (izquierda) y aspecto típico
del macho con el plastron cóncavo y la cola larga (derecha), fotos J.M. Renjifo, A. González
150 • Tortugas y Crocodylia
 Fused vertebrae
Ribs
Carapace

Neck

REPTILES: Testudinos
to de este grupo de organismos,
idad del grupo, en virtud de que
a y no el ataque, como se deduce
reducidas son bastante agresivas
- Figure 28-6
tes ambientes; así por ejemplo se
milia Testudinidae poseen grandes
eados o en forma de domo, como
ontra los • Presencia de oído medio e interno
predadores, a la vez que
• Poca audición
exposición directa a los rayos del
ncos o exponiendo el dorso del
• Buen olfato
cesos digestivos, crecer y madurar
ar algas y otros ectoparásitos. Por
uícolas tienen conchas aplanadas
nto debajo Buena visión
• del agua.
• Mandíbulas sin dientes tles are nonetheless a varied
• Ovíparas
Las tortugas son vertebrados amniotas, anápsidos y ectotérmicos (las
cuales regulan la temperatura corporal mediante un activo intercambio
con el medio ambiente), muy antiguos, se originaron en el Triásico me-
dio y son los tetrápodos Plastronmás altamente especializados en su morfología.
Estos animales longevos, con tasas de fertilidad reducidas, periodos de
Skeleton and shell of a turtle, maduración
showing fusionsexual tardíosand
of vertebrae y tasas de crecimiento
ribs with the carapace.muy lentas,
The long andtienen
flexibleíndices
neck allows the turtle to
withdraw its head into its shellde
for reclutamiento
protection. muy bajos y resultan por ende bastante sensibles a la
sobreexplotación y a la destrucción de sus hábitats.
two layers: an outer horny layer of ker-
Las costillas de las tortugas son únicas entre todos los vertebrados
atin and an inner layer of bone. New
tetrápodos puesto que se encuentran fusionadas a la concha (no son
layers of keratin are laid down beneath
móviles), lo cual afecta el proceso respiratorio, dado que se tiene que
the old as the turtle grows and ages.
The bony layer is a fusion resolver
of ribs,elverte-
problema de inhalar y exhalar dentro de una armadura rígida.
Como todosele-
brae, and many dermally-ossifying los vertebrados tetrápodos las tortugas respiran a través de
pulmones,
ments (Figure 28-6). Unique amongsin embargo varias de ellas pueden absorber oxígeno directa-
ver-
tebrates, turtle limbs and
are located inside theirrigadas
mentelimb delgirdles
agua por diversos métodos, bien sea a través de zonas altamente
del epitelio bucal, la faringe o la piel. De todos estos mecanismos
ribs! Lacking
teeth, the turtle jaw is provided with
tough, horny plates forcloaca receptáculo profusamente vascularizado denominado bursa
en unfood
(Figure 28-7).
gripping
cloacal, hacia donde el individuo bombea agua regularmente para atrapar
Clumsy and unlikelyelasoxígeno disuelto; esta forma de respiración es utilizada para prolongar
they appear
to be within their protective los periodos de permanencia subacuática y rehuir predadores o para
shells, tur-
logically diverse group that
hibernar andbajo el agua.
eco-
Lasseems ablecarecen de dientes, en su
tortugas Figure 28-7
to adjust to human presence.
lugar poseen un estuche córneo
Snapping turtle,llamado
Chelydra serpentina, showing the absence of teeth. Instead, the jaw edges are
The terms “turtle,” “tortoise,”
ranfoteca, que recubre
and “terrapin”
las mandíbulas
covered with a horny de
forma similar al pico de las aves, con el cual
plate.

are applied variously to different members

trituran y desgarran the
of the turtle order. In North American
los alimentos.
limbs andPese the rigid shell. The sating for poor hearing are a goo
• Nidos en tierra usage, they are all correctlyacalled
lo que se podría suponer
turtles. movements los órganos
of the limbs during walk- sense of smell, acute vision, and col
de losgiven
The term “tortoise” is frequently sentidos
to se encuentran
ing also help bastante
to ventilate the lungs. perception evidently as good as that
desarrollados
land turtles, especially the large forms. en esteMany grupoaquatic
de reptiles; la gain enough oxy-
turtles humans.
British usage of the terms isvisión de las
different: tortugasgen
“tor- es muyby aguda ya que water in and out
just pumping Turtles are oviparous. Fertilizatio
• Sexo depende de la temperatura
toise” is the inclusive term,pueden
is applied only to the aquatic
whereaspercibir
“turtle” movimientos extraños a mouth
of a vascularized cavity; this
Mandíbulas en forma de
is internal and all turtles, even marin
gancho
unamembers.
distancia mínimaactivity
de 100 enables
m. La retina them to remain sub- forms, bury their shelled, amniotic egg
del nido
las tortugas (1-marinas, 2-terrestres, contiene más conos que merged for long
bastones, es decir
de un tapaculo (Kinosternon sp.)
periods when inac- in the ground. Usually considerab
(1, 3) y en forma de domo (2) One consequence células of living in a
fotosensibles tive. When
adaptadas paraactive
la visiónthey must
diurna y enlung- care is exercised in constructing th
color. El olfato
 Squamata: fusion of snout Archosauria: presence of
bones, characteristics of opening anterior to eye,
palate, skull roof, vertebrae, orbit shaped like inverted

REPTILES: Squamata
ribs, pectoral girdle, humerus

Lepidosaurians: character-
istics of bone growth,
triangle, teeth laterally
compressed

skull, pelvis, feet

• Grupo reciente Dorsal
temporal
d-roofed anapsid opening Orbit
and carapace
• 95% de las especies de
ermal bone and
reptiles vivientes: lagartos y
axial skeleton Lateral
temporal
serpientesDiapsids: diapsid skull opening
with 2 pairs of temporal
openings
• Cráneo díapsido y ventana Diapsid skull

palatina
Turtle-diapsid clade (Sauropsida) Figure 28-2
characteristics of skull
Cladogram of the living Amniota showing monophyletic groups. Some of th
and appendages
shared derived characters (synapomorphies) that are diagnostic for the
• Dos
: extraembryonic
caracteres: serpientes lineages are given. The skulls represent the ancestral condition of the three
groups. The skulls of modern diapsids and synapsids are often highly mod
es of amnion,
by loss or fusion of skull bones that obscures the ancestral condition.
nd allantois
• Elongación del cuerpo vs Representative skulls for anapsids are Nyctiphruetus of the upper Permian;
rearreglo de los órganos diapsids, Youngina of the upper Permian; for synapsids, Aerosaurus, a
pelycosaur of the lower Permian. The relationships expressed in this cladog
are tentative and controversial, especially that between birds and mammals
• Consumo de grandes presas Contrary to the view shown here, in which mammals are the outgroup, som
authorities support a sister-group relationship between birds and mammals