Chapter 26 - Angelo State University

”
Amniotic “Pond
Pond”
Chapter 26
ƒ Any animal with a shell-less egg remains tied
to water.
ƒ The development of a shelled egg freed the
reptilian groups to exploit land.
ƒ Extraembryonic membranes from previous
evolutionary aquatic stages are maintained.
ƒ Surrounding the whole young organism is a
porous, parchment-like or leathery shell.
ƒ
Amniote Origins
and
Reptilian Groups
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Amniotes
ƒ The Amniota is a
single, monophyletic
lineage of Paleozoic
tetrapods with this
developmental pattern.
ƒ Before the end of the
Paleozoic Era,
amniotes diverged into
reptilian groups, birds
and mammals.
Diversity
ƒ Members of the paraphyletic class Reptilia
include the first truly terrestrial vertebrates.
ƒ Nearly 7000 species are described; about 300
occur in the United States and Canada.
ƒ The Age of Reptiles lasted over 165 million
years and included the dinosaurs.
ƒ A mass extinction occurred at the end of the
Mesozoic; modern reptiles represent surviving
lineages.
History
ƒ Amniotes arose from amphibian-like tetrapods, the anthracosaurs,
during the Carboniferous.
ƒ By the late Carboniferous, amniotes had separated into three
lineages.
– Anapsids have a skull with no temporal opening behind the orbits; modern
turtles are anaspids.
– Diapsids gave rise to all other reptilian groups and to the birds.
• The diapsid skull has two temporal openings; one pair below the cheeks
and another above.
• Lepidosaurs include ichthyosaurs and modern reptiles except for turtles
and crocodilians.
• More derived archosaurs included dinosaurs, living crocodilians, and birds.
• Sauropterygians included extinct aquatic groups including the long-necked
plesiosaurs.
– Synapsids are mammal-like reptiles with a single pair of temporal openings.
Fig. 26.2
1
Changes in Traditional Classification of Reptilian Groups
Classification
ƒ Cladistic methodology insists on hierarchical
arrangement of monophyletic groups.
ƒ This disqualifies the traditional class Reptilia as a
valid taxon because it is not monophyletic.
ƒ Class Reptilia excludes birds, which descend
from the most recent common ancestor of
reptiles.
ƒ This makes the class Reptilia a paraphyletic
group because it does not include all descendants
and their most recent common ancestor.
ƒ Reptiles are therefore identified as amniotes that are not birds.
ƒ Based solely on shared derived characteristics, crocodilians and
birds are sister groups belonging to a monophyletic group apart
from other reptiles, the Archosauria.
ƒ Some taxonomists defined Reptilia as the Archosauria plus the
lepidosaurs, thus including birds.
ƒ Evolutionary taxonomists argue that birds represent a novel
adaptive zone and grade of organization; they maintain the class
Aves based on the morphological and ecological novelty of birds.
ƒ Reptilian group” refers to members of four monophyletic groups
formerly considered class Reptilia
Are Birds living Dinosaurs?
Distinguishing Reptiles from Amphibians
ƒ Reptiles and birds share
several derived characters,
including several skull
characteristics and a largely
aglandular skin with a
special type of harder
keratin called Beta keratin,
which unites them as a
monophyletic group.
Scales
ƒ Reptile scales are primarily of keratin, formed from
epidermis, and not homologous with fish scales.
ƒ Scales may grow gradually to replace wear, as in alligators.
ƒ Snakes and lizards replace old scales with new and “shed their
skins.”
Skin
ƒ Reptiles have a tough, dry, scaly skin that
offers protection against desiccation and
injury.
ƒ The thin epidermis is shed periodically.
ƒ A thicker, well-developed dermis underneath
has chromatophores that provide color.
Other Skin Adaptations
ƒ Turtles add new layers underneath old layers
of platelike scutes.
ƒ Crocodiles and many lizards have bony plates
called osteoderms underneath the keratinized
scale.
ƒ Resistance to desiccation is provided by
hydrophobic lipids in the epidermis, as well as
the presence of Beta keratin.
2
Amniotes and the Amniotic Egg
ƒ The amniotic egg contains four extraembryonic
membranes including the amnion.
– Embryos develop within the amnion, a membranous sac that
provides an aquatic environment and protects it from shock.
– The yolk sac is the first membrane and pre-dates the amniotes
by millions of years
– The allantois is a sac that grows out of the hindgut and holds
metabolic wastes during development.
– The chorion lies beneath the eggshell and encloses the rest of
the embryo.
ƒ With growth, the allantois and chorion fuse to form the
chorioallantoic membrane, a provisional “lung.”
Shelled Egg of Reptiles
ƒ A shelled egg contains food and protective
membranes to support embryonic development
on land.
ƒ Reptiles lay eggs on land; young hatch as lungbreathing juveniles, not as aquatic larvae.
ƒ The shelled egg widened the division between
evolving amphibians and reptiles.
ƒ More than any other adaptation, this
contributed to the evolutionary establishment
of reptiles.
Internal Fertilization
ƒ A shelled egg requires internal fertilization; the
sperm must reach the egg before it is enclosed.
ƒ Paired testes produce sperm that are carried by
vasa deferentia to the copulatory organ.
ƒ The female has paired ovaries and oviducts.
ƒ Glandular walls of the oviducts secrete albumin
and shells for the relatively large eggs.
Fig. 26.4
Circulatory System Modifications
ƒ Reptiles have an efficient circulatory system and higher blood
pressure than amphibians.
ƒ The right atrium that receives unoxygenated blood is completely
partitioned from the left atrium.
ƒ Crocodilians have separated ventricles dividing pulmonary and
systemic circulation.
ƒ Other reptiles have an incompletely separated ventricle, but little
mixture of blood occurs and the effect is two functionally separate
circulations.
ƒ The incomplete separation between the right and left sides of the
heart in some groups provides an added benefit of permitting
blood to bypass the lungs when pulmonary respiration is not
occurring (e.g., during diving or aestivation).
Lungs
ƒ Reptile lungs are better developed than those
of amphibians.
ƒ Reptiles depend almost exclusively on lungs,
supplemented by pharyngeal membranes.
ƒ Reptiles pull air into the lungs by enlarging the
thoracic cavity.
ƒ Snakes and lizards use expansion of the rib
cage; turtles and crocodilians use movement of
internal organs to create negative pressure to
inhale.
3
Feeding
ƒ Reptilian Jaws
– The jaws of fish
allowed fast jaw
closure to seize food
but little force for
chewing.
– In reptiles, jaw
muscles became
larger and arranged
for the mechanics of
chewing.
Support of Limbs for Locomotion
Water Conservation
ƒ All amniotes have a metanephric kidney drained by
the ureter.
ƒ Nephrons of the reptilian metanephros lack the loop
of Henle that allows concentration of solutes.
ƒ Many reptiles have salt glands near the nose or eyes
to secrete salty fluid hyperosmotic to body fluids.
ƒ Nitrogenous wastes are excreted as uric acid rather
than urea or ammonia.
ƒ The amniotic egg of reptiles permits rapid
development of large young in relatively dry
environments.
Fig. 26.5
ƒ Except for limbless members, all reptiles have
better body support than amphibians.
ƒ Many modern reptiles still walk with legs
splayed outward and the belly close to the
ground.
ƒ Most dinosaurs and some modern lizards have
more efficient legs directed beneath the body.
Nervous System
ƒ The reptilian nervous system is considerably more
complex than the amphibian.
ƒ The reptile brain is still small but the cerebrum is
relatively larger.
ƒ Crocodilians have the first true cerebral cortex; it
allows complex behaviors not seen in amphibians.
ƒ Sense organs are generally well developed, except for
hearing.
ƒ The Jacobson’s organ is highly developed in lizards
and snakes to detect odors carried by the tongue
Natural History of Reptilian Orders
Anapsid Reptiles: Subclass Anapsida
ƒ Order Testudines includes the turtles.
ƒ Turtles descended from one of the earliest
anapsid lineages, probably in the late Permian.
ƒ Fossils appear in the Upper Triassic, 200 million
years ago, and they have occurred ever since.
ƒ Turtles lack teeth and have tough, horny plates
for gripping food.
4
Shells
ƒ The shells consist of the dorsal carapace and the ventral
plastron.
ƒ The shell has an outer horny layer of keratin and an inner
layer of bone.
ƒ The bony layer is a fusion of ribs, vertebrae, and dermallyossified elements.
ƒ Unique among vertebrates, the limbs and limb girdles are
located deep to the ribs.
Nervous System and Senses
ƒ The turtle brain is small, less than one percent of its
body weight.
ƒ The cerebrum is larger than that of amphibians; some
turtles can learn a path through a maze.
ƒ Turtles have a middle and an inner ear but sound
perception is poor; they make few sounds aside from
those made during mating.
ƒ Turtles have a good sense of smell, acute vision, and
color perception about equal to humans.
Order Squamata
Squamata:: Lizards, Snakes and Worm Lizards
ƒ Squamates are the most recent and diverse of
diaspids; they make up 95% of living reptiles.
ƒ Lizards appeared in the fossil record in the
Permian but did not radiate until the
Cretaceous.
ƒ Snakes appeared in the late Cretaceous from a
group whose descendants include monitor
lizards.
ƒ Snakes gained specializations for losing their
legs and therefore for engulfing large prey.
Breathing
ƒ One consequence of having a rigid shell is that the turtle cannot
expand its chest to breathe.
ƒ Turtles solve this problem by using abdominal and pectoral
muscles as a “diaphragm.”
ƒ Air is drawn in by contraction of the limb flank muscles, increasing
abdominal cavity volume.
ƒ Exhalation is accomplished by drawing back the shoulder girdle to
compress the viscera.
ƒ These actions are visible as bellows-like movements at the turtle’s
“limb pockets.”
ƒ Movement of limbs during walking also helps ventilate the lungs.
ƒ Some aquatic turtles gain sufficient oxygen by pumping water in
and out of the mouth cavity.
Reproduction and Developmen
Developmentt
ƒ Turtles are oviparous; fertilization is internal
and all turtles bury their eggs in the ground.
ƒ Once the female lays her eggs in a nest, she
deserts them.
ƒ In some turtle families, as in crocodilians and
some lizards, nest temperature determines the
sex of hatchlings; low temperatures produce
males and high temperatures produce females.
ƒ Diaspid skulls have lost dermal
bone ventral and posterior to the
lower temporal opening.
ƒ This allowed evolution in lizards of
a mobile skull with movable joints,
a kinetic skull.
ƒ The quadrate, fused to the skull in
other reptiles, has a joint at the
dorsal end and articulates with the
lower jaw.
ƒ Joints in the palate and across the
roof of the skull allow the snout to
be tilted up.
ƒ This allows squamates to seize and
manipulate prey, and effectively
close the jaw with force.
ƒ Exceptional skull mobility of
snakes is considered a major factor
in their diversification.
Skull
Modifications
5
Viviparity
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Viviparity in reptiles is limited to squamates.
It has evolved at least 100 separate times.
It is associated with cold climates.
Viviparity involves increasing the length of
time eggs are kept in the oviduct.
ƒ Developing young respire through
extraembryonic membranes.
ƒ Young obtain nutrition from yolk sacs or via
the mother, or a combination of both.
Suborder Sauria
Sauria:: Lizards
ƒ Diverse group with terrestrial, burrowing, aquatic, arboreal, and some
aerial members.
ƒ Some have degenerate limbs; the glass lizards are nearly limbless.
ƒ Have movable eyelids whereas snakes have a transparent covering.
ƒ Nocturnal geckos only rods; day-active lizards have both rods and
cones.
ƒ Have an external ear that snakes lack.
ƒ Some survive well in hot and dry regions.
– Conserve water by producing semisolid urine with high content of crystalline uric acid.
– Some can store fat in their tails to provide energy and metabolic water during drought.
ƒ The Gila monster and beaded lizard are only lizards capable of a
venomous bite.
ƒ Lizards keep their body temperature relatively constant by behavioral
thermoregulation, although they are ectotherms.
ƒ Ectothermy is a successful strategy in ecosystems with low
productivity and warm climates, such as tropical deserts and
grasslands.
Suborder Serpentes
Serpentes:: Snakes
ƒ Snakes are limbless and have lost the pectoral and pelvic
girdles (except in pythons).
ƒ The many vertebrae are shorter and wider than in other
tetrapods, allowing undulation.
ƒ Elevation of the neural spine gives the musculature more
leverage.
ƒ The feeding apparatus allows them to eat prey several times
their own diameter.
ƒ The two halves of the lower jaw are loosely joined, allowing
them to spread apart.
ƒ The skull bones also are loosely articulated so the mouth can
accommodate a large prey.
ƒ To keep breathing during the slow process of swallowing, the
tracheal opening is extended.
Snake Features
ƒ Eyeballs have reduced mobility and a permanent corneal
membrane for protection.
ƒ Most snakes have poor vision, but arboreal snakes in tropical
forests have highly developed vision.
ƒ Snakes lack external ears and do not respond to most aerial
sounds.
ƒ Snakes can feel vibrations at low frequencies, especially vibrations
carried in the ground.
ƒ Chemical senses rather than vision or hearing are the main senses
snakes use to hunt prey.
ƒ Jacobson’s organs are a pair of pits in the roof of the mouth; they
are lined with olfactory epithelium and the forked tongue picks up
scent particles and conveys them past this organ.
ƒ A snake’s skin is infolded between scales; when stretched by a big
meal, the skin is unfolded.
Fig. 26.18a
Snake Locomotion
ƒ Lateral undulation is
the S-shaped movement
that pushes against
rough ground and water.
ƒ Concertina movement
is extension of S-shaped
loops to strike or to
climb trees.
6
Snake Locomotion
Pit vipers
ƒ Have “pits” with nerve endings
sensitive to heat emitted by warmbodied birds and mammals.
ƒ Viper fangs are hollow and hinged
to inject venom when the snake
strikes.
ƒ Of an average of 8000 snake bites
each year in the U.S., only about
12 result in death.
ƒ Nonvenomous snakes kill prey by
constriction or by biting and
swallowing.
ƒ Rectilinear movement
is a straight movement
using minute lifting of
consecutive ribs.
ƒ Sidewinding is a
sideways looping by
desert vipers that
“walks” them across
loose sand.
Snakebite and Toxicity
ƒ Saliva of harmless snakes contains limited toxins;
this provided a basis for natural selection of venom.
ƒ Most snake venoms are a complex combination of
venom types.
ƒ Neurotoxins act on the nervous system, causing
blindness or stopping respiration.
ƒ Hemorrhagin type venoms broke down blood vessels;
much blood is leaked into tissue spaces.
ƒ Toxicity is measured by the median lethal dose on
laboratory animals, called the LD50.
ƒ Sea snakes and the Australian tiger snake have the
most deadly venom per unit.
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Large venomous snakes deliver more
venom; a king cobra may be the most
dangerous.
In India and Burma, dense populations
and poor footwear contribute to 200,000
bites per year.
There are about 8,000 snakebites per year
in the United States, resulting in only
about 12 deaths.
The world total for deaths from snakebite
is about 50,000 to 60,000 each year, with
most deaths occurring in India, Pakistan,
Myanmar, and nearby countries where
poorly shod people frequently come into
contact with venomous snakes or do not
get immediate medical attention once bit.
Less than 20 percent of all snakes are
venomous, although venomous species
outnumber nonvenomous species by 4 to
1 in Australia.
Snakebite and
Toxicity
Source
Source
ƒ Most snakes are oviparous
and lay shelled eggs under
logs or rocks or in holes in
the ground.
ƒ Others, including pit
vipers, are ovoviviparous.
ƒ A few snakes are
viviparous, having a
primitive placenta to
exchange nutrients with
the young.
ƒ Snakes can store sperm
and lay several clutches of
fertile eggs long after a
single mating.
Reproduction
7
Order Crocodilia
Crocodilia:: Crocodiles and Alligators
ƒ Modern crocodilians are the only surviving reptiles
of the archosaurian lineage.
ƒ This lineage gave rise to the Mesozoic radiation of
dinosaurs and to birds.
ƒ Modern crocodilians differ little from primitive
crocodilians of the early Mesozoic.
ƒ All have a long, well-reinforced skull and jaw
musculature for a powerful bite.
ƒ A type of dentition called thecodont, where teeth are
set in sockets, was typical of archosaurs.
ƒ They have a complete secondary palate, a feature
only shared with mammals.
Crocodilians
Source
Source
Reproduction
ƒ Alligators and crocodiles are oviparous;
usually 20-50 eggs are laid in a mass of
vegetation.
ƒ Nests left unguarded are easily discovered and
raided by predators.
ƒ High nest temperatures produce males; low
temperatures produce females and can result in
females outnumbering males five to one.
The End
8