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• Diapsidos Pm
Single opening for
external nares
Incisors Incisive
foramen
Canine

• Amniotas, diversos M L
Premolars

N Molars Palatal
foramina
F

• Cocodrilos parientes cercanos J

J
Pal
Pt
Internal
nares
V
Foramen
Prs ovale
Carotid

Huevos con cáscara

Bs
•
canal
P Sq
Sq Bo Glenoid
Posterior
lacerate Middle
foramen ear
Occipital
Occipital

Tipo de huevo y clivaje

(a) (b)

•
condyles

N
P F
L Pm FIGURE 7.51
Sq J M of the opossum D
palatal (b), and later

• Estructuras óseas y Mastoid

Pt Molars Premolars
Incisors
Abbreviations: basio
basisphenoid (Bs), d
(F), jugal (J), lacrima

musculatura Middle
(N), parietal (P), pa
D Canine
ear (Pm), presphenoid
Inflected angle squamosal (Sq).
(c) of dentary After Carroll.

• Primera vertebral cervical Interparietal

Orbitosphenoid

encaja con el condilo

Sq
Presphenoid
Alisphenoid
Supraoccipital

occipital
Exoccipital
Foramen Lingula Tympan
magnum
Basioccipital Basisphenoid Pterygoid P
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hic09617_ch29.qxd 6/7/00 2:14 PM Page 583
• Un hueso auditibo, estribo
• Desechos de nitrógeno en
forma de ácido úrico
• Dinosaurios
A B
• Estructura de la pelvis, Figure 29-1
Archaeopteryx, a 147-million-year-old ancestor of modern birds. A, Cast of the second and most nearly perfect fossil of Archaeopteryx, which was discovered in
a Bavarian stone quarry. Seven specimens of Archaeopteryx have been discovered, the most recent one in 1992. B, Reconstruction of Archaeopteryx.
clavícula y muñeca disappointingly meager. The finding
was also dramatic because it proved
beyond reasonable doubt the phylo-
genetic relatedness of birds and
reptiles.
Registros fósiles
Zoologists had long recognized the
• similarity of birds and reptiles. The
skulls of birds and reptiles abut against
the first neck vertebra by a single ball-
and-socket joint, the occipital condyle
(mammals have two condyles). Birds
and reptiles have a single middle ear
bone, the stapes (mammals have three
middle ear bones). Birds and reptiles
have a lower jaw composed of five or
six bones, whereas the lower jaw of
mammals has one mandibular bone, the
dentary. Birds and reptiles excrete their
nitrogenous wastes as uric acid whereas
mammals excrete theirs as urea. Birds
and reptiles lay similar yolked eggs with
the early embryo developing on the sur-
face by shallow cleavage divisions.
The distinguished English zoolo-
gist Thomas Henry Huxley was so
impressed with these and many other
anatomical and physiological affinites
that he called birds “glorified reptiles”
and classified them with a group of
dinosaurs called theropods that dis-
played several birdlike characteristics
(Figures 29-2 and 29-3). Theropod
dinosaurs share many derived charac-
ters with birds, the most obvious of
which is the elongate, mobile, S-
shaped neck. As shown in the clado-
gram (Figure 29-3), theropods belong
to a lineage of diapsid reptiles, the
archosaurians, that includes crocodil-
ians and pterosaurs, as well as the
dinosaurs. There is now overwhelming
evidence that Huxley was correct:
birds’ closest phylogenetic affinity is to
the theropod dinosaurs. The only
anatomical feature required to link bird
ancestry with the theropod dinosaurs
CHAPTER 29 Birds 583
was feathers, and this was provided by
the discovery of Archaeopteryx. Recent
discoveries of Cretaceous bird fossils in
Spain, Madagascar, and China provide
new data on bird ancestry. All these
new discoveries, however, still link
early birds with theropod reptiles.
Living birds (Neornithes) are
divided into two groups: (1) Paleog-
nathae (Gr. palaios, ancient, !
gnathos, jaw), the large flightless
ostrichlike birds and the kiwis, often
called ratite birds, which have a flat
sternum with poorly developed pec-
toral muscles, and (2) Neognathae (Gr.
neos, new, ! gnathos, jaw), flying birds
that have a keeled sternum on which
powerful flight muscles insert. This divi-
sion originated from the view that
flightless birds (ostrich, emu, kiwi, rhea)
represented a separate line of descent
that never attained flight. This idea is
now completely rejected. Ostrichlike
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Melanin
granules

• Escamas de reptiles elaboradas Epidermal

cell

Chromatophore
• Escamas epidérmicas en patas
kar24239_ch06_212-239.qxd 12/22/10 3:43 PM Page 222 (in epidermis)

y pies
Chr

• Presencia de vasos sanguíneos, (a)

terminaciones nerviosas y Melanin

FIGURE 6.14 Bird scales Stratum and skin. (a) Epidermal scales ar
corneum

musculo liso (Dermis, cerca granules Transitional layer layer, the stratum corn
the stratum basale and the keratinized surface
stratum intermedium and transitional layer before reaching the surfac
del foliculo plumoso) Epidermal
cell
Stratum intermedium
granulosum layers of mammals. Epidermis

(a) After Smith; (b) after Lucas and Stettenheim.

Chromatophore Stratum basale
• Dermis del pecho, muy (in epidermis)
Dermis
Bird skin has few glands. The uropygial gland, located
vascularizada formando un at the base of the tail (figure 6.15a), secretes a lipid and pro-
Chromatophore
“terreno incubador” tein product that birdsBlood capillary
collect on the sides of their beak and
then smear on their feathers. Preening coats the feathers with
(a) (b)
this secretion, making them water repellent, and probably
FIGURE 6.14 Bird scales and skin. (a) Epidermal scales conditions theon
are present keratin
the feetofand
which they
legs of are(b)composed.
birds. Follow-
Section of skin showing
ing a molt, preening also helps the newly regenerated feather
the stratum basale and the keratinized surface layer, the stratum corneum. Cells moving out of the basal layer move through first the
stratum intermedium and transitional layer before reaching theunfurl
surface.and
Theseassume its functional
middle layers shape.
are equivalent to the The otherand
spinosum gland,
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• Epidermis: estrato basal y

córneo.
Melanin Stratum corneum

• Capa de células en transición granules Transitional layer

que se transforman en la Epidermal

cell
Stratum intermedium Epidermis

superficie queratinizada del Chromatophore Stratum basale

estrato córneo (in epidermis)

Dermis

Chromatophore Blood capillary

• Piel con pocas glándulas
(a) (b)

• Gl. Uropigial, producto

FIGURE 6.14 Bird scales and skin. (a) Epidermal scales are present on the feet and legs of birds. (b) Section of skin showin
the stratum basale and the keratinized surface layer, the stratum corneum. Cells moving out of the basal layer move through first the
lipoproteico
stratum intermedium and transitional layer before reaching the surface. These middle layers are equivalent to the spinosum and
granulosum layers of mammals.
(a) After Smith; (b) after Lucas and Stettenheim.

• Gl. Sal, aves marinas

Bird skin has few glands. The uropygial gland, located Feathers develop embryologically from feather f
at the base of the tail (figure 6.15a), secretes a lipid and pro- cles, invaginations of the epidermis that dip into the un
tein product that birds collect on the sides of their beak and lying dermis. The root of the feather follicle, in associa
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• Plumas, derivados de la piel
Pulp

Sin vasos sanguíneos ni

caps
• shed

nervios Feather Sheath

filament kar24239_ch06_212-239.qxd 12/22/10 slough
3:43 PM Page 223

• Pterilas Dermal
Skin
surface
core
Feather
follicle

• Foliculo plumoso, Dermal

papilla

invaginación epidérmica
(a) (b) (c) (d)

FIGURE 6.16 Feather growth. Molting and developmental sequence of feather replacement. (a) The old feather is
and a new feather filament soon grows out of the follicle as a result of cell proliferation at its base. (b, c) Successive stages in
development. Note that some tissues necessary for initial development (sheath, pulp caps) now lose this function and are slou
the mature feather emerges. (d) Mature, new feather in place.

• Raíz del foliculo plumoso vs Based on the research of P. F. A. Maderson and W. J. Hillenius

morphogenetic signals emanating from the patterning zone.

Pterylae

not only sets cell fates but also presides over

pulpa dérmica, forman la Here cells become programmed to form sheath, pulp caps,
barbs, barbules, or rachis. Cells moving through the patterning
zone receive different signals than cells that precede or follow
pluma them, leading to the highly specific differentiation of the
emerging feather. As the spathe is being differentiated, the pat-
terning process sets aside populations of keratinocytes, for
Ventral
example, tissues of the future barb, barbules, and rachis. Addi-
tionally, other signals also establish precisely patterned fates
(a)
where cells will lose their connections to one another and form
the future spaces and slits between barbs and barbules. Thus
between feather parts, and also programs cells des
the sheath, pulp caps, and possibly the stratum cy
well as the feather primordium itself. The rachis i
by the fusion of several barbs but also by this patter
Uropygial
gland
Functions There are several types of feathers (f
ContourDorsal
feathers aerodynamically shape the su
bird. Down feathers lie close to the skin as the
tion. Filoplumes are often specialized for d
(b) Feath
flight feathers constitute the major aerodynam
 Rachis
Pterylae

Down
Interlocking feather
barbules

AVES: TEGUMENTO
Uropygial
gland
Contour Barb
Ventral Dorsal feather

(a)
(b) Feather types Calamus

• Funciones: Vane
Barb

Rachis

• Plumas de contorno, forma Contour

feather
FIGURE 6.15 Feather tracts and
feather morphology. (a) Feathers arise

aerodinámica along specific pterylae or feather tracts.

(b) General morphology of contour and
down feathers. (c) Feathers types. Flight
feathers constitute the major locomotor
surfaces. Contour feathers on the body

Aislante térmico
aerodynamically shape the surface of a bird.
• Calamus (quill)
Down feather Filoplume Filoplumes are often specialized for display.
Down feathers lie close to the skin as
Flight feather thermal insulation
(c) Feather types (a, c) After Smith; (b) after Spearman.

• Filoplumas, plumas de exhibición

• Plumas de vuelo
• Remeras, raquis largo o
prominente (locomoción)
above it that will generate the morphogenetic signals pre-
siding over the fates of these keratinocytes. In the follicle,
rings of outer cells (beta-keratin), the sheath and feather
itself, form more or less concentrically around the inner
stratum cylindricum and pulp caps (alpha-keratin), and der-
mal core (figure 6.17, cross sections). The feather filament
continues to grow out from the follicle accompanied by the
highly vascularized dermal core, which extends through the
follicle mouth above the surrounding integument. Core
tissues are protected from desiccation and trauma by a suc-
cession of pulp caps derived from the stratum cylindricum.
The protective sheath, important initially as a scaffold
to the developing feather, is eventually lost to preening
once the differentiated feather is mature and ready to
unfurl. The spathe is the first part of the young feather to
differentiate beneath the sheath. As the tip of the spathe
unfurls, the base of the spathe is still under construction.
When the spathe completes its differentiation, the calamus
is next formed, also in the same region beneath the sheath.
As calamus formation proceeds, pulp caps continue to form
within its hollow core as the dermal core regresses within
the follicle. Dermal muscles, connected in a network of
muscles, act to erect mobile feathers.
The patterning process is complex. New keratinocytes
formed in the proliferation zone move up in the follicle
but their fates are determined in the patterning zone by

Integument 223

• Estimulos sensoriales y colores

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Nasofrontal hinge

• Caja craneana dilatada y

osificada
(a)
• Sin dientes, estuches Quadrate Pterygoid Palatine Jugal
bar
Upper
jaw

queratinizados

• Mandíbulas alargadas, pico

• Aves neognathas, paladar (b) Orbital septum

(parasphenoid)

dividido

• Articulación pterigo-
palatina, cráneo
procinetico
(c)
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Nasofrontal hinge
• Caja craneana dilatada y
osificada
Premaxilla

• Sin dientes, estuches

Quadrate
queratinizados
Pterygoid Palatine Jugal Upper
bar jaw
Maxilla
• Mandíbulas alargadas, pico
Palatine
• Aves paleonagthas, paladar Jugal bar
no unido Pterygoid

Basipterygoid
Orbital septum
• Se desliza a través de
(parasphenoid)
process
Quadrate
salientes (d)
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(teleosts)
fication o

• Columna vertebral flexible Ver

stome fi
depends u
Cervical ers propu
vertebrae produce l
• Movilidad de la cabeza the ossifi
as the ma
Int
Synsacrum
composed
• Vertebras, parte media y between
that mak

posterior de la columna such a w

within th
proteins
special ch
ing water
• Fusionadas con la cintura vice of re
In t
pelvica marily as
body duri
into later
lations of
• Rigidez y estabilidad, early basi
ment, the

condiciones para el vuelo suspendin

buoyancy
tetrapod
design th
such as br
• Menos músculos para el FIGURE 8.41 Bird vertebral column. Regions of
extensive vertebral fusion are indicated posteriorly within the
about or
becomes
control vertebral synsacrum.The numerous heterocoelous cervical vertebrae allow
for great mobility of the head.
appearan
brae, such
neural sp
the verte