6. Overview: Life Without a Backbone
• Invertebrates are animals that lack a backbone
• They account for 95% of known animal species
• They are morphologically diverse
– For example, the Christmas tree worm
(Spirobranchus giganteus) has tentacles for gas
exchange and feeding
7. Figure 33.3a
Porifera (5,500 species) Placozoa (1 species)
0.5 mm
A sponge
Cnidaria (10,000 species)
A placozoan (LM)
Ctenophora (100 species)
A jelly
Acoela (400 species)
1.5 mm
Acoel flatworms (LM) A ctenophore, or comb jelly
9. Figure 33.3c
Loricifera (10 species) Priapula (16 species) Onychophora (110 species)
50 µm
A loriciferan (LM) A priapulan
An onychophoran
Nematoda Tardigrada Arthropoda
(25,000 species) (800 species) (1,000,000 species)
100 µm
A roundworm
(colored SEM)
Tardigrades A scorpion (an arachnid)
Ecdysozoa (colorized SEM)
10. Figure 33.3d
Hemichordata Chordata
(85 species) (52,000 species)
A tunicate
Echinodermata (7,000 species)
An acorn worm
Deuterostomia
A sea urchin
11. Sponges are basal animals that lack true
tissues
• Animals in the phylum Porifera are known
informally as sponges
• They are sedentary and live in marine waters or
fresh water
13. • Sponges are suspension feeders, capturing food
particles suspended in the water that passes
through their body
• Water is drawn through pores into a cavity called
the spongocoel and out through an opening
called the osculum
• Sponges lack true tissues and organs
14. Figure 33.4
Food particles
in mucus
Flagellum Choanocyte
Azure vase sponge Choanocyte Collar
(Callyspongia
plicifera) Osculum
Spongocoel Phagocytosis of
food particles
Amoebocyte
Pore
Spicules
Epidermis
Water
flow
Amoebocytes
Mesohyl
17. Figure 33.4c
Food particles
in mucus
Flagellum
Choanocyte
Collar
Phagocytosis of
food particles
Amoebocyte
18. • Choanocytes, flagellated collar cells, generate a
water current through the sponge and ingest
suspended food
• Sponges consist of a gelatinous noncellular
mesohyl layer between two cell layers
• Amoebocytes are found in the mesohyl and play
roles in digestion and structure
• Most sponges are hermaphrodites: Each
individual functions as both male and female
19. Cnidarians are an ancient phylum of
eumetazoans
• All animals except sponges and a few other
groups belong to the clade Eumetazoa, animals
with true tissues
• Phylum Cnidaria is one of the oldest groups in this
clade
21. • Cnidarians have diversified into a wide range of
both sessile and motile forms including jellies,
corals, and hydras
• They exhibit a relatively simple diploblastic, radial
body plan
• The basic body plan of a cnidarian is a sac with a
central digestive compartment, the
gastrovascular cavity
• A single opening functions as mouth and anus
22. • There are two variations on the body plan: the
sessile polyp and motile medusa
• A polyp adheres to the substrate by the aboral
end of its body
• A medusa has a bell-shaped body with its mouth
on the underside
• Medusae do not attach to the substate but move
freely
24. • Cnidarians are carnivores that use tentacles to
capture prey
• The tentacles are armed with cnidocytes, unique
cells that function in defense and capture of prey
• Nematocysts are specialized organelles within
cnidocytes that eject a stinging thread
28. Hydrozoans
• Most hydrozoans alternate between polyp and
medusa forms
• Hydra, a freshwater cnidarian, exists only in polyp
form and reproduces asexually by budding
29. Figure 33.8-1
Reproductive
Feeding polyp
polyp
Medusa
bud Gonad
Medusa
Portion
of a
colony
of polyps
Key
1 mm
Haploid (n)
Diploid (2n)
30. Figure 33.8-2
Reproductive
Feeding polyp
polyp
Medusa MEIOSIS
bud Gonad
Medusa
Egg Sperm
SEXUAL
REPRODUCTION
Portion
of a
colony FERTILIZATION
of polyps
Zygote
Key
1 mm
Haploid (n)
Diploid (2n)
31. Figure 33.8-3
Reproductive
Feeding polyp
polyp
Medusa MEIOSIS
bud Gonad
Medusa
Egg Sperm
SEXUAL
ASEXUAL
REPRODUCTION
Portion REPRODUCTION
of a (BUDDING)
colony FERTILIZATION
of polyps
Zygote
Developing
Planula
polyp
(larva)
Mature
polyp Key
1 mm
Haploid (n)
Diploid (2n)
32. Scyphozoans
• In the class Scyphozoa, jellies (medusae) are the
prevalent form of the life cycle
33. Cubozoans
• In the class Cubozoa, which includes box jellies
and sea wasps, the medusa is box-shaped and
has complex eyes
• Cubozoans often have highly toxic cnidocytes
34. Anthozoans
• Class Anthozoa includes the corals and sea
anemones, and these cnidarians occur only as
polyps
• Corals often form symbioses with algae and
secrete a hard external skeleton
35. Lophotrochozoans, a clade identified by
molecular data, have the widest range of
animal body forms
• Bilaterian animals have bilateral symmetry and
triploblastic development
• Most have a coelom and a digestive tract with two
openings
• The clade Bilateria contains Lophotrochozoa,
Ecdysozoa, and Deuterostomia
37. • The clade Lophotrochozoa was identified by
molecular data
• Some develop a lophophore for feeding, others
pass through a trochophore larval stage, and a
few have neither feature
• Lophotrochozoa includes the flatworms, rotifers,
ectoprocts, brachiopods, molluscs, and annelids
38. Flatworms
• Members of phylum Platyhelminthes live in
marine, freshwater, and damp terrestrial habitats
• Although flatworms undergo triploblastic
development, they are acoelomates
• They are flattened dorsoventrally and have a
gastrovascular cavity with one opening
• Gas exchange takes place across the surface, and
protonephridia regulate the osmotic balance
39. • Flatworms are divided into two lineages
– Catenulida, or “chain worms,” reproduce asexually
by budding
– Rhabditophora are more diverse and include both
free-living and parasitic species
41. Free-Living Species
• The best-known rhabditophorans are planarians
• Planarians live in fresh water and prey on smaller
animals
• Planarians have light-sensitive eyespots and
centralized nerve nets
42. • The planarian nervous system is more complex
and centralized than the nerve nets of cnidarians
• Planarians are hermaphrodites and can reproduce
sexually, or asexually through fission
44. Parasitic Species
• Parasitic rhabditophorans live in or on other
animals
• Two important groups of parasitic rhabditophorans
are the trematodes and the tapeworms
45. Trematodes
• Trematodes parasitize a wide range of hosts, and
most have complex life cycles with alternating
sexual and asexual stages
• Trematodes that parasitize humans spend part of
their lives in snail hosts
• They produce surface proteins that mimic their
host and release molecules that manipulate the
host’s immune system
46. Figure 33.11
Male
Human
host
Female
1 mm
Mature flukes
Motile Ciliated
larva larva
Snail host
47. Tapeworms
• Tapeworms are parasites of vertebrates and
lack a digestive system
• Tapeworms absorb nutrients from the host’s
intestine
48. • The scolex contains suckers and hooks for
attaching to the host
• Proglottids are units that contain sex organs and
form a ribbon behind the scolex
• Fertilized eggs, produced by sexual reproduction,
leave the host’s body in feces
50. Rotifers
• Rotifers, phylum Rotifera, are tiny animals that
inhabit fresh water, the ocean, and damp soil
• Rotifers are smaller than many protists but are
truly multicellular and have specialized organ
systems
51. Figure 33.13
Jaws Crown
of cilia
Anus
Stomach 0.1 mm
52. • Rotifers have an alimentary canal, a digestive
tube with a separate mouth and anus that lies
within a fluid-filled pseudocoelom
• Rotifers reproduce by parthenogenesis, in which
females produce offspring from unfertilized eggs
• Some species are unusual in that they lack males
entirely
53. Lophophorates: Ectoprocts and Brachiopods
• Lophophorates have a lophophore, a crown of
ciliated tentacles around their mouth
• Lophophorates have a true coelom
• Lophophorates include two phyla: Ectoprocta and
Brachiopoda
54. • Ectoprocts (also called bryozoans) are sessile
colonial animals that superficially resemble plants
• A hard exoskeleton encases the colony, and
some species are reef builders
56. • Brachiopods superficially resemble clams and
other hinge-shelled molluscs, but the two halves of
the shell are dorsal and ventral rather than lateral
as in clams
• Brachiopods are marine and attach to the seafloor
by a stalk
57. Molluscs
• Phylum Mollusca includes snails and slugs,
oysters and clams, and octopuses and squids
• Most molluscs are marine, though some inhabit
fresh water and some snails and slugs are
terrestrial
• Molluscs are soft-bodied animals, but most are
protected by a hard shell
58. • All molluscs have a similar body plan with three
main parts
– Muscular foot
– Visceral mass
– Mantle
• Many molluscs also have a water-filled mantle
cavity and feed using a rasplike radula
60. • Most molluscs have separate sexes with gonads
located in the visceral mass, but many snails are
hermaphrodites
• The life cycle of many molluscs includes a ciliated
larval stage called a trochophore
61. • Four of the major classes of molluscs are
– Polyplacophora (chitons)
– Gastropoda (snails and slugs)
– Bivalvia (clams, oysters, and other bivalves)
– Cephalopoda (squids, octopuses, cuttlefish, and
chambered nautiluses)
62. Chitons
• Chitons are oval-shaped marine animals encased
in an armor of eight dorsal plates
• They use their foot like a suction cup to grip rock,
and their radula to scrape algae off the rock
surface
68. • Most gastropods are marine, but many are
freshwater and terrestrial species
• The most distinctive characteristic of gastropods is
torsion, which causes the animal’s anus and
mantle to end up above its head; torsion is
different from the coiling of a shell
• Most have a single, spiraled shell
• Slugs lack a shell or have a reduced shell
69. Figure 33.18
Mantle Stomach Intestine
cavity
Anus
Mouth
70. Bivalves
• Bivalves are marine and include many species of
clams, oysters, mussels, and scallops
• They have a shell divided into two halves drawn
together by adductor muscles
• Some bivalves have eyes and sensory tentacles
along the edge of their mantle
72. • The mantle cavity of a bivalve contains gills that
are used for feeding as well as gas exchange
73. Figure 33.20
Coelom
Hinge area
Mantle Gut Heart Adductor muscle
(one or two)
Digestive
gland
Anus
Mouth
Excurrent
siphon
Shell
Palp Water
flow
Foot
Mantle Incurrent
cavity Gonad Gill siphon
74. Cephalopods
• Cephalopods are carnivores with beak-like jaws
surrounded by tentacles of their modified foot
• Most octopuses creep along the sea floor in
search of prey
76. • Squids use their siphon to fire a jet of water, which
allows them to swim very quickly
77. • One small group of shelled cephalopods, the
nautiluses, survives today
78. • Cephalopods have a closed circulatory system,
well-developed sense organs, and a complex
brain
• Shelled cephalopods called ammonites were
common but went extinct at the end of the
Cretaceous 65.5 million years ago
79. Protecting Freshwater and Terrestrial
Molluscs
• Molluscs are the animal group with the largest
number of recent extinctions
• The most threatened groups are
– Freshwater bivalves, including pearl mussels
– Terrestrial gastropods, including Pacific island
land snails
• These molluscs are threatened by habitat loss,
pollution, and non-native species
80. Figure 33.22
Molluscs
Other invertebrates Amphibians
Insects
Birds
Fishes
Mammals Reptiles
(excluding
An endangered Pacific Recorded extinctions of animal
birds)
island land snail, species (Data: International Union for
Partula suturalis Conservation of Nature, 2008)
Workers on a mound of pearl
mussels killed to make buttons
(ca. 1919)
81. Figure 33.22b
Molluscs
Other invertebrates Amphibians
Insects
Birds
Fishes
Mammals Reptiles
(excluding
Recorded extinctions of animal birds)
species (Data: International Union for
Conservation of Nature, 2008)
82. Figure 33.22c
Workers on a mound of pearl
mussels killed to make buttons
(ca. 1919)
83. Annelids
• Annelids have bodies composed of a series of
fused rings
• Annelids are coelomates
• The phylum Annelida is divided into two groups
– Polychaeta (polychaetes)
– Oligochaeta (earthworms and their relatives, and
leeches)
84. Polychaetes
• Members of class Polychaetes have paddle-like
parapodia that work as gills and aid in locomotion
• Most polychaetes are marine
86. Oligochaetes
• Oligochaetes (class Oligochaeta) are named for
relatively sparse chaetae, bristles made of chitin
87. Earthworms
• Earthworms eat through soil, extracting nutrients
as the soil moves through the alimentary canal
• Earthworms are hermaphrodites but cross-fertilize
• Some reproduce asexually by fragmentation
90. Leeches
• Most species of leeches live in fresh water; some
are marine or terrestrial
• Leeches include predators of invertebrates, and
parasites that suck blood
• Leeches secrete a chemical called hirudin to
prevent blood from coagulating
92. Ecdysozoans are the most species-rich
animal group
• Ecdysozoans are covered by a tough coat called a
cuticle
• The cuticle is shed or molted through a process
called ecdysis
• The two largest phyla are nematodes and
arthropods
94. Nematodes
• Nematodes, or roundworms, are found in most
aquatic habitats, in the soil, in moist tissues of
plants, and in body fluids and tissues of animals
• They have an alimentary canal, but lack a
circulatory system
• Reproduction in nematodes is usually sexual, by
internal fertilization
96. • Caenorhabditis elegans is a model organism in
research
• Some species of nematodes are important
parasites of plants and animals
• Thrichinella spiralis can be acquired by humans
from undercooked pork
98. Arthropods
• Two out of every three known species of animals
are arthropods
• Members of the phylum Arthropoda are found in
nearly all habitats of the biosphere
99. Arthropod Origins
• The arthropod body plan consists of a segmented
body, hard exoskeleton, and jointed appendages
• This body plan dates to the Cambrian explosion
(535–525 million years ago)
• Early arthropods show little variation from segment
to segment
101. • Arthropod evolution is characterized by a
decrease in the number of segments and an
increase in appendage specialization
• These changes may have been caused by
changes in Hox gene sequence or regulation
102. Figure 33.29
EXPERIMENT
Other
ecdysozoans
Origin of Ubx and
abd-A Hox genes?
Arthropods
Common ancestor Onychophorans
RESULTS
Ant = antenna
J = jaws
L1–L15 = body segments
103. General Characteristics of Arthropods
• The appendages of some living arthropods are
modified for functions such as walking, feeding,
sensory reception, reproduction, and defense
104. Figure 33.30
Cephalothorax Abdomen
Antennae Thorax
Head
(sensory
reception)
Swimming appen-
dages (one pair per
abdominal segment
Walking legs
Pincer Mouthparts
(defense) (feeding)
105. • The body of an arthropod is completely covered by
the cuticle, an exoskeleton made of layers of
protein and the polysaccharide chitin
• When an arthropod grows, it molts its exoskeleton
106. • Arthropods have eyes, olfactory receptors, and
antennae that function in touch and smell
• Arthropods have an open circulatory system in
which hemolymph is circulated into the spaces
surrounding the tissues and organs
• A variety of organs specialized for gas exchange
have evolved in arthropods
107. • Molecular evidence suggests that living arthropods
consist of four major lineages that diverged early
in the phylum’s evolution
– Chelicerates (sea spiders, horseshoe crabs,
scorpions, ticks, mites, and spiders)
– Myriapods (centipedes and millipedes)
– Hexapods (insects and relatives)
– Crustaceans (crabs, lobsters, shrimps,
barnacles, and many others)
108. Chelicerates
• Chelicerates, subphylum Chelicerata, are named
for clawlike feeding appendages called chelicerae
• The earliest cheliceriforms were eurypterids
(water scorpions)
• Most marine cheliceriforms (including eurypterids)
are extinct, but some species survive today,
including horseshoe crabs
112. • Arachnids have an abdomen and a cephalothorax,
which has six pairs of appendages: the chelicerae,
the pedipalps, and four pairs of walking legs
• Gas exchange in spiders occurs in respiratory
organs called book lungs
• Many spiders produce silk, a liquid protein, from
specialized abdominal glands
113. Figure 33.33
Intestine Stomach
Heart Brain
Digestive
gland
Eyes
Ovary
Poison
gland
Anus
Book lung
Spinnerets Gonopore Chelicera Pedipalp
(exit for eggs) Sperm
Silk gland receptacle
114. Myriapods
• Subphylum Myriapoda includes millipedes and
centipedes
• Myriapods are terrestrial, and have jaw-like
mandibles
• Millipedes eat decaying leaves and plant matter
• Millipedes have many legs, with two pairs per
trunk segment
116. • Centipedes are carnivores
• Centipedes have one pair of legs per trunk
segment
117. Insects
• Subphylum Hexapoda, insects and relatives, has
more species than all other forms of life combined
• They live in almost every terrestrial habitat and in
fresh water
• The internal anatomy of an insect includes several
complex organ systems
119. • Insects diversified several times following the
evolution of flight, adaptation to feeding on
gymnosperms, and the expansion of angiosperms
• Insect and plant diversity declined during the
Cretaceous extinction, but has been increasing in
the 65 million years since
120. • Flight is one key to the great success of insects
• An animal that can fly can escape predators, find
food, and disperse to new habitats much faster
than organisms that can only crawl
122. • Many insects undergo metamorphosis during their
development
• In incomplete metamorphosis, the young, called
nymphs, resemble adults but are smaller and go
through a series of molts until they reach full size
123. • Insects with complete metamorphosis have
larval stages known by such names as maggot,
grub, or caterpillar
• The larval stage looks entirely different from the
adult stage
125. • Most insects have separate males and females
and reproduce sexually
• Individuals find and recognize members of their
own species by bright colors, sound, or odors
• Some insects are beneficial as pollinators, while
others are harmful as carriers of diseases, or
pests of crops
• Insects are classified into more than 30 orders
127. Crustaceans
• While arachnids and insects thrive on land,
crustaceans, for the most part, have remained in
marine and freshwater environments
• Crustaceans, subphylum Crustacea, typically have
branched appendages that are extensively
specialized for feeding and locomotion
• Small crustaceans exchange gases through the
cuticle; larger crustaceans have gills
128. • Most crustaceans have separate males and
females
• Isopods include terrestrial, freshwater, and marine
species
– Pill bugs are a well known group of terrestrial
isopods
• Decapods are all relatively large crustaceans and
include lobsters, crabs, crayfish, and shrimp
134. Echinoderms and chordates are
deuterostomes
• Echinoderms (phylum Echinodermata) include sea
stars and sea urchins
• Chordates (phylum Chordata) include the
vertebrates
• Echinoderms and chordates constitute the clade
Deuterostomia
135. • Deuterostomes share developmental
characteristics
– Radial cleavage
– Formation of the anus from the blastopore
• However, some other animals also share these
developmental characteristics
• Deuterostomes are defined primarily by DNA
similarities
137. Echinoderms
• Sea stars and most other echinoderms are
slow-moving or sessile marine animals
• A thin epidermis covers an endoskeleton of hard
calcareous plates
• Echinoderms have a unique water vascular
system, a network of hydraulic canals branching
into tube feet that function in locomotion and
feeding
• Males and females are usually separate, and
sexual reproduction is external
138. Figure 33.40
Short digestive tract
Stomach
Anus
Spine
Gills
Central disk
Madreporite
Radial
nerve
Digestive glands
Ring Gonads
canal
Ampulla
Podium
Radial canal Tube feet
139. • Most adult echinoderms have radial symmetry with
multiples of five
• Echinoderm larvae have bilateral symmetry
140. • Living echinoderms are divided into five classes
– Asteroidea (sea stars and sea daisies)
– Ophiuroidea (brittle stars)
– Echinoidea (sea urchins and sand dollars)
– Crinoidea (sea lilies and feather stars)
– Holothuroidea (sea cucumbers)
141. Asteroidea: Sea Stars and Sea Daisies
• Sea stars have multiple arms radiating from a
central disk
• The undersurface of each arm bears tube feet,
which grip substrate with adhesive chemicals
• Sea stars feed on bivalves by prying them open
with their tube feet, everting their stomach, and
digesting their prey externally with digestive
enzymes
• Sea stars can regrow lost arms
142. • Sea daisies were discovered in 1986, and only
three species are known
• Sea daisies live on submerged wood and absorb
nutrients through a membrane that surrounds their
body
144. Ophiuroidea: Brittle Stars
• Brittle stars have a distinct central disk and long,
flexible arms, which they use for movement
• Some species are suspension feeders, while
others are predators or scavengers
146. Echinoidea: Sea Urchins and Sand Dollars
• Sea urchins and sand dollars have no arms but
have five rows of tube feet
• Their spines are used for locomotion and
protection
• Sea urchins feed on seaweed using a jaw-like
structure on their underside
148. Crinoidea: Sea Lilies and Feather Stars
• Sea lilies live attached to the substrate by a stalk
• Feather stars can crawl using long, flexible arms
• Both are suspension feeders
150. Holothuroidea: Sea Cucumbers
• Sea cucumbers lack spines, have a very reduced
endoskeleton, and do not look much like other
echinoderms
• Sea cucumbers have five rows of tube feet; some
of these are developed as feeding tentacles
152. Chordates
• Phylum Chordata consists of two subphyla of
invertebrates as well as hagfishes and vertebrates
• Chordates are bilaterally symmetrical coelomates
with segmented bodies
• Chordates share many features of embryonic
development with echinoderms, but have evolved
separately for at least 500 million years