Lecture Notes

to accompany
Life: The Science of Biology, Eighth Edition
Sadava • Heller • Orians • Purves • Hillis

Chapter 31: Animal Origins and the Evolution of Body

Plans

Introduction
• Evolutionary biologists are using developmental and genetic knowledge, combined with fossil evidence, to
understand how a few fundamental body plans could have been modified to yield a remarkable diversity of
animal forms.

What Evidence Indicates the Animals Are

Monophyletic?
• What traits distinguish the animals from the other groups of organisms?
• In contrast to the Bacteria, Archaea, and most microbial eukaryotes, all animals are multicellular.
• In contrast to plants, all animals are heterotrophs.
• In contrast to fungi, animals use internal processes to break down materials from their environment.
• In contrast to plants, animals can move.
Animal monophyly is supported by gene sequences and morphology
• Molecular and morphological evidence indicates that all animals are descendants of a single ancestral
lineage.
• All animals share a derived set of traits:
• Similarities in their ribosomal RNAs.
• Similarities in their Hox genes.
• Special types of cell–cell junctions: tight junctions, desmosomes, and gap junctions.
• A common set of extracellular matrix molecules, including collagen.
• Animals evolved from ancestral colonial flagellated protists.
• Within these ancestral colonies of cells, functional specialization arose.
• Different cells became specialized for different functions, such as movement and nutrition, and others
differentiated into gametes.
• Once the division of labor had begun, the specialized units continued to differentiate while improving
their coordination with other working groups of cells.
• These coordinated groups of cells evolved into animals.
• Clues to the evolutionary relationships among animal groups must be sought in derived traits that are found
in some groups but not in others.
• Such characters can be found in fossils, in patterns of embryonic development, in the morphology and
physiology of living animals, in the structure of molecules of animals, and in the genomes of animals.
Developmental patterns show evolutionary relationships among animals
• Figure 31.1 shows a current phylogenetic tree of animals.
• Developmental patterns of embryonic development, which have provided important clues about animal
phylogeny, are more evolutionarily labile than previously thought.
• Cleavage is the first few cell divisions of a zygote.
• Different cleavage patterns exist among animals.
 • Cleavage patterns may be complete or incomplete and are influenced by the configuration of the yolk.
(See Section 43.1.)
• Radial cleavage is when the egg cell divides in an even pattern. This is the ancestral condition for
eumetazoans and is found among many protostomes and diploblastic animals as well as deuterostomes.
• Spiral cleavage is a complicated derived form of radial cleavage. Spiral cleavage is found among many
lophotrochozoans, which are sometimes known as spiralians.
• Early branches of ecdysozoans have radial cleavage, but most ecdysozoans have a cleavage pattern that
is neither radial nor spiral in organization.
• Animals form layers of cells during their development from a single-celled zygote to a multicellular adult.
• The embryos of diploblastic animals have two cell layers: an outer ectoderm and an inner endoderm.
• The embryos of triploblastic animals have a third layer, the mesoderm, which lies between the ectoderm
and the endoderm.
• The existence of three cell layers distinguishes the protostomes and deuterostomes from simple animals
that diverged earlier.
• Protostomes and deuterostomes differ in the fate of the blastopore that forms in the spherical embryo.
• In the protostomes, the mouth arises from the blastopore. (See Figure 31.2.)
• In the deuterostomes, the blastopore gives rise to the anus.
• The process of gastrulation is central to our understanding of evolutionary relationships among the animals.

What Are the Features of Animal Body Plans?

• The entire structure of an animal, its organ systems, and the integrated functioning of its parts are known as
its body plan.
• Animal body plans can be seen as variations on four key features: Symmetry, structure of the body cavity,
segmentation, and external appendages that move the body.
Most animals are symmetrical
• The overall shape of an animal’s body plan is referred to as its symmetry.
• A symmetrical animal can be divided along at least one plane into similar halves.
• Animals that have no plane of symmetry are said to be asymmetrical.
• There are different types of symmetry. (See Figure 31.3.)
• Spherical symmetry is the simplest type of symmetry, in which body parts radiate out from a central point.
Spherical symmetry is widespread among the protists.
• An organism with radial symmetry has one main axis around which its body parts are arranged. (See Figure
31.3A.)
• Animals that exhibit radial symmetry include some simple sponges, sea anemones, and most Cnidarians
and Ctenophores.
• Bilaterally symmetric animals can be divided into mirror images by a single plane that passes through the
midline of their bodies from the front to the back end. (See Figure 31.3B.)
• Bilateral symmetry is a common characteristic of animals that move freely through their environments.
• Bilateral symmetry is often associated with cephalization: the presence of a head bearing sensory organs
and central nervous tissues at the anterior end of the animal.
The structure of the body cavity influences movement
• Body cavities are fluid-filled spaces that lie between the cell layers of the bodies of many kinds of animals.
• The type of body cavity an animal has influences how it can move.
• Acoelomate animals lack an enclosed, fluid-filled cavity. (See Figure 31.4A.)
• These animals move by beating cilia.
• Body cavities come in two types, and they are differentiated by their relationship to the mesoderm.
• Pseudocoelomate animals have a body cavity called a pseudocoel, a fluid-filled space in which many of
the internal organs are suspended. There is no mesoderm surrounding the internal organs. (See Figure
31.4B.)
• Coelomate animals have a coelom, a body cavity that develops within the mesoderm. The coelom is
enclosed on both the inside and outside by mesoderm. (See Figure 31.4C.)
• The fluid-filled body cavities of simple animals function as hydrostatic skeletons.
 • When the muscles surrounding fluids contract, the fluids can be moved to other flexible parts of the
body, causing these body regions to expand.
• Most animals also have hard skeletons that provide protection and facilitate movement.
Segmentation improves control of movement
• Segmentation facilitates specialization of different body regions and allows an animal to alter the shape of
its body in complex ways, and to control movements precisely.
• Segmentation evolved independently in both the protostomes and the deuterostomes.
• Segmentation is variable. In some animals, segments are not unapparent externally, in others they are
repeated many times, or are visible but differ strikingly. (See Figure 31.5.)
Appendages enhance locomotion
• Locomotion in animals is important for finding food, avoiding predators, and finding mates.
• External appendages enhance an animal’s ability to move around.
• Many echinoderms have tube feet.
• Arthropods and vertebrates have jointed limbs.
• Jointed limbs have been a prominent factor in the evolutionary success of both of these groups.
• In arthropod insects, the pterosaurs, the birds, and the bats, body plans emerged in which limbs were
modified into wings.

How Do Animals Get Their Food?

• Most animals must actively obtain food.
• Sensory structures that can provide animals with detailed information about their environment, as well as
nervous systems that can receive, process, and coordinate that information have been favored by evolution.
• Motile animals move from one place to another.
• Sessile animals stay in one place.
• There are a few broad categories of animal feeding strategies:
• Filter feeders capture small organisms delivered to them by their environment.
• Herbivores eat plants or parts of plants.
• Predators capture and eat other animals.
• Parasites live in or on other organisms.
• Detritivores feed on dead organic material.
• A variety of feeding strategies can be found among all the major animal groups.
Filter feeders capture small prey
• Filter feeders use some kind of straining device to filter food from the air or water environment.
• Many sessile aquatic animals rely on water currents to bring prey to them. (See Figure 31.6.)
• Motile filter feeders bring the nutrient-containing medium to them.
• Some sessile filter feeders expend energy to move water past their food-capturing devices.
• Choanocytes are specialized flagellated feeding cells in sponges.
Herbivores eat plants
• Many different kinds of herbivores may feed on a single kind of plant, consuming different parts of the
plant or eating the same part in different ways. (See Figure 31.8.)
• Herbivores often feed on plants without killing them.
• Herbivores typically have long complex guts designed to digest tough or fibrous plant tissues. Plants tissues
may also contain chemicals that must be detoxified before they can be ingested.
Predators capture and subdue large prey
• Predators possess features, such as sensitive sensory organs, sharp teeth, and behavioral adaptations, that
enable them to find, capture, and subdue relatively large prey. (See Figure 31.9.)
• Cnidarians (jellyfish and their relatives) use specialized cells that contain stinging organelles called
nematocysts, to capture and subdue prey that are much larger and more complex then themselves. (See
Figure 31.10.)
• Omnivores are animals that eat both plants and other animals.
 • The diet of many animals differs at different life stages.
Parasites live in or on other organisms
• Parasites are animals that live in or on another organism— called a host—and obtain their nutrients by
consuming parts of that organism.
• Parasites are often much smaller than their hosts and consume parts of their host without killing it.
• Parasites have complex life cycles and often rely on many hosts.
• Endoparasites are morphologically simple organisms that live inside their hosts.
• Ectoparasites are more complex and have digestive tracts and mouthparts that enable them to pierce a
host’s tissues or suck out their body fluids.

How Do Animal Life Cycles Differ?

• The life cycle of an animal encompasses its embryonic development, birth, growth to maturity, reproduction,
and death.
• Direct development refers to animals that have newborns that are similar to adults.
• A larva is an immature life cycle stage that has a form different from that of an adult.
• Life cycles with metamorphosis have striking life cycle changes. (See Figure 31.11.)
• In these animals, different stages may be specialized for different functions, which may increase the
efficiency with which the animal performs particular tasks such as feeding and reproducing.
All life cycles have at least one dispersal stage
• Dispersal refers to when an animal moves, or is moved, so that it does not die exactly where it was born.
• Animals that are sessile as adults typically disperse eggs or larvae.
• Trochophore larvae have a distinctive form and are found in several marine animal clades with spiral
cleavage, most notably polychaete worms and the mollusks. (See Figure 31.12A.)
• Nauplius are bilaterally symmetrical larvae that will mature into a crustacean with a segmented body and
jointed appendages. (See Figure 31.12B.)
• Animals that are motile as adults, such as moths and butterflies, typically disperse once they are mature.
• Some animals disperse during several different life cycle stages.
No life cycle can maximize all benefits
• The characteristics of an animal in any one life cycle stage may improve its performance in one activity,
but reduce its performance in another—a situation known as a trade-off.
• Some major trade-offs can be seen in animal reproduction.
• With a fixed amount of available energy, a female animal can produce many small eggs or a few large
eggs, but she cannot do both. (See Figure 31.13.)
• Incubation times or eggs vary between species of birds. (See Figure 31.14.)
• Altricial young hatch when they are young and helpless. Parents can provide for a small number of
altricial young.
• Precocial young hatch when they are developed to a point that they are able to forage for themselves
almost immediately. Parents of precocial young must incubate their eggs longer.
Parasite life cycles evolve to facilitate dispersal and overcome host defenses
• Animals that live as internal parasites may not need to exert much energy to obtain food, but they must
overcome their host’s defenses, and either they or their offspring must disperse to new hosts while their
host is still living.
• Most parasite species have complex life cycles involving one or more intermediate hosts and several larval
stages. (See Figure 31.15.)
• Complex life cycles may facilitate the transfer of individual parasites among hosts.

What Are the Major Groups of Animals?

• Table 31.1 provides a summary of the living members of the major animals groups.
• The Bilateria is a large monophyletic group embracing all animals other than sponges, ctenophores, and
cnidarians.
 • Members of the Bilateria have bilateral symmetry, three cell layers, and the presence of at least seven Hox
genes.
• Bilaterian animals are classified as either protostomes or deuterostomes.
• Sponges are not a clade, but the name is used for three groups that exhibit ancestral body organization of
animals.
• Eumetazoans are all other animals, including the bilaterians.
• Eumetazoans have obvious body symmetry, a gut, a nervous system, special types of cell junctions, and
well-organized tissues in distinct cell layers.
Sponges are loosely organized animals
• Sponges are the simplest animals.
• They have some specialized cells, but no distinct layers, no true organs, and no body symmetry.
• They have hard skeletal elements called spicules.
• There appear to be three major groups of sponges. (See Figure 31.16.)
• Glass sponges and demosponges have skeletons made of silicaceous spicules; these are composed of
hydrated silicon dioxide.
• Calcareous sponges have calcium carbonate skeletons.
• The body plan of sponges is an aggregation of cells built around a water canal system.
• Specialized feeding cells called choanocytes line the inside of the water canal. (See Figures 31.7 and
31.16A.)
• Sponges have a supporting skeleton of simple or branching spicules and often have an elastic network of
fibers.
• Most sponges are filter feeders; a few are carnivores.
• Most of the 8,000 sponge species are marine.
• Sponges depend on water movement through their bodies to obtain food and are often oriented at right
angles to current flow so that they may intercept water and the prey it contains as it flows past them.
• Sponges reproduce both sexually and asexually.
• In most species, a single individual produces both eggs and sperm.
• Asexual reproduction is by budding and fragmentation.
Ctenophores are radially symmetrical and diploblastic
• Ctenophores (or comb jellies) lack most Hox genes possessed by all other eumetazoans.
• They are radially symmetrical and diploblastic with two cell layers separated by gelatinous mesoglea.
• Ctenophores, unlike cnidarians, have a complete gut; food enters through a mouth, and wastes are voided
through two anal pores.
• Ctenophores have eight comblike rows of fused plates of cilia, called ctenes, which they use for movement.
• Ctenophore tentacles do not have nematocysts; rather, they are covered with sticky filaments to which prey
adhere.
• The ctenophores also have low metabolic rates.
• All 100 known species live in the open sea.
• Their life cycles are uncomplicated: Gametes from gonads located on the walls of the gastrovascular cavity
are released into the cavity and then discharged through the mouth or pores.
• Fertilization takes place in the open seawater, and the fertilized egg develops into a miniature ctenophore
that grows into an adult.
• (See Video 31.3.)
Cnidarians are specialized carnivores
• The mouth of a cnidarian is connected to a blind sac called the gastrovascular cavity. It functions in
digestion, circulation, gas exchange, and acts as a hydrostatic skeleton.
• This single opening serves as both mouth and anus.
• The generalized cnidarian life cycle has two stages: the sessile polyp and the motile medusa. (See Figure
31.18.)
• The polyp is typically asexual; individual polyps may reproduce by budding to form colonies.
• The medusae produce eggs and sperm and release them into the water.
 • When an egg is fertilized, it forms a free-swimming, ciliated larva called a planula that eventually settles to
the bottom and transforms into a polyp.
• The polyp and the medusa stages share a similar body plan; a medusa is essentially a polyp without a stalk.
• Cnidarians have epithelial cells with muscle fibers whose contractions allow them to move, as well as nerve
nets that integrate body activities.
• They have specialized structural molecules (collagen, actin, and myosin) and Hox genes.
• Cnidocytes have structures called nematocysts that can discharge toxins into their prey. (See Figure 31.10.)
• The cnidarian body plan combines a low metabolic rate with the ability to capture large prey, allowing
cnidarians to survive in environments where prey is scarce.
• There are about 11,000 species living today. (See Figure 31.19.)
• Scyphozoans:
• The several hundred species of scyphozoans are all marine.
• They are commonly known as jellyfish or sea jellies.
• The medusa typically has the form of an inverted cup, and the tentacles with nematocysts extend
downward from the margin of the cup.
• The medusa dominates the life cycle of the scyphozoans. (See Figure 31.18.)
• Anthozoans:
• Members of the anthozoan clade are all widespread in warm and cold ocean waters.
• The anthozoans include sea anemones, sea pens, and corals.
• The anthozoans entirely lack the medusa stage of the life cycle.
• Polyps produce eggs and sperm, and the fertilized egg develops into a planula that develops directly into
more polyps.
• Sea anemones are solitary. Some species are able to swim, others can crawl slowly, and some can
burrow. (See Figure 31.19A.)
• The sea pens are generally sessile and colonial. (See Figure 31.19B.)
• Corals are also usually sessile and colonial. (See Figure 31.20.)
• The polyps of corals secrete a matrix of organic molecules on which calcium carbonate is deposited.
• This matrix forms the eventual skeleton of the coral colony.
• As coral colonies grow, old polyps die and leave their calcareous skeletons behind. (See Figure
31.20B.)
• Living members of the colony form a layer on top of a growing reef of skeletal remains.
• Coral formations more than 2,000 km long and 150 km wide make up the Great Barrier Reef along the
northeastern coast of Australia.
• Corals flourish in nutrient-poor, clear, tropical waters.
• Symbiotic, photosynthetic dinoflagellates help corals obtain enough nutrients to grow rapidly.
• The corals provide protection for the dinoflagellates which provide the products of photosynthesis to
the corals.
• Both global warming and nutrient runoff are threatening coral reefs throughout the world.
• Hydrozoans:
• Life cycles are diverse among the hydrozoans.
• The polyp commonly dominates the life cycle, but some species have only medusae, whereas others have
only polyps.
• Figure 31.21 illustrates the life cycle of colonial hydrozoan polyps.
• (See Videos 31.1 and 31.2, and Animated Tutorial 31.1.)

Supplemental Photo Collection

The Instructor’s Media Library includes photographs of animals from many of the major and minor taxa discussed in
the textbook, including sponges, cnidarians, and ctenophores.