Trilobites, Chelicerates, and

Myriapods
Chapter 19
Phylum Arthropoda

l Two out of every

three known
species of animals
are arthropods.
l Members of the
phylum
Arthropoda are
found in nearly all
habitats of the
biosphere.
Phylum Arthropoda

l Arthropods are:
l Multicellular
l Bilaterally symmetrical

l Triploblastic

l Have a true coelom (protostomes)

l Segmented
General Characteristics of
Arthropods
l The diversity and success of arthropods
are largely related to their segmentation,
hard exoskeleton (made of chitin), and
jointed appendages.
General Characteristics of
Arthropods
l Segments have combined into functional
groups called tagmata.
l Tagmata have specialized purposes.
General Characteristics of
Arthropods
l Arthropods have an open circulatory
system in which fluid called hemolymph
is circulated into the spaces surrounding
the tissues and organs.
General Characteristics of
Arthropods

Why have arthropods

achieved such great
diversity and
abundance?
A Versatile Exoskeleton
l The exoskeleton of
arthropods is very
protective, but still
flexible.
l The exoskeleton is
made of chitin.
l Prevents desiccation.
l Provides places for
muscle attachment.
l Does not allow for
growth, the outer
covering must be
molted – ecdysis.
More Efficient Locomotion
l Usually, each
segment bears a pair
of jointed
appendages.
l The appendages
have sensory hairs
and may be modified
for sensory functions,
food handling, or
walking & swimming.
Air Piped Directly to Cells
l Most terrestrial
arthropods have an
efficient tracheal
system of air tubes,
which delivers oxygen
directly to the tissues
and cells.
l Limits body size.
l Aquatic arthropods
breathe using internal or
external gills.
Highly Developed Sensory Organs

l Arthropods have a
variety of sensory
organs.
Complex Behavior Patterns

l Arthropods show
complex behavior
patterns.
l Mostly innate
behaviors.
l Some learned.
Metamorphosis

l Intraspecific
competition (between
members of one
species) is reduced
because of
metamorphosis.
l Larval forms may be
quite different from
adults.
Relationships Among Arthropod
Subgroups
l Clade Panarthropoda,
Phylum Arthropoda
l Divided into subphyla based
on relationships between
subgroups based on
molecular data.
l Centipedes, millipedes,
pauropods, and symphylans
are placed in subphylum
Myriapoda.
l Insects are placed in
subphylum Hexapoda.
l Spiders, ticks, horseshoe
crabs and their relatives form
subphylum Chelicerata.
l Lobsters, crabs, barnacles,
and others form subphylum
Crustacea.
Relationships Among Arthropod
Subgroups
l Formerly, insects and
myriapods were placed
together in uniramia.
l Uniramous
appendages.
l The “mandibulate
hypothesis” suggests all
arthropods with
mandibles are more
closely related to each
other than to arthropods
with chelicerae.
l Mandibles in each
group may or may not
be homologous.
Subphylum Trilobita

l Early arthropods, such as trilobites showed

little variation from segment to segment.
Subphylum Trilobita

l Trilobites arose
during the Cambrian
– maybe earlier and
lasted for 300 million
years.
Subphylum Trilobita

l Trilobites had a trilobed shape.

l Three tagmata:
l Head (cephalon) with a mouth, compound
eyes, antennae, and 4 pairs of leglike
appendages.
l Trunk with a variable number of segments
each with a pair of biramous appendages.
l One of the branches of biramous appendage
was fringed and may have been a gill.
l Pygidium – segments fused into a plate.
Subphylum Trilobita

l Most could roll up like pill bugs.

l Probably benthic scavengers.
l Many (especially later species) had large,
complex, many-faceted eyes.
Subphylum Chelicerata

l Chelicerate arthropods include

eurypterids, horseshoe crabs, spiders,
ticks, mites, scorpions, & sea spiders.
Subphylum Chelicerata
l They have 6 pairs
of
cephalothoracic
appendages:
l Chelicerae
(mouthparts)
l Pedipalps
l 4 pairs of
walking legs
l Lack mandibles
and antennae.
Class Merostomata

l Class Merostomata
includes the eurypterids
and horseshoe crabs.
l Eurypterids were giant
water scorpions up to 3
m in length.
l Cambrian through
Permian.
l Predators, some with
large crushing claws.
Class Merostomata

l Three genera of
horseshoe crabs
live today.
l Limulus, found in
North America,
has existed on
earth almost
unchanged since
the Triassic period.
Class Merostomata

l Horseshoe crabs have an unsegmented

carapace (hard dorsal shield), a broad
abdomen, and a long telson (tail piece).
l Cephalothorax
l Chelicerae
l Pedipalps
l 4 pairs walking legs
l Abdomen
l 6 pairs of thin appendages
l Book gills found on 5.
Class Merostomata
l Horseshoe crabs have simple and compound eyes.
l Feed at night on worms and small molluscs.
l Come to shore in large numbers to mate at high tide.
l Trilobite larvae resemble trilobites.
Class Pycnogonida

l Sea spiders, class

Pycnogonida, have
small, thin bodies and
usually 4 pairs of
walking legs.
l Found in all oceans,
most common in polar
seas.
l Some have chelicerae
and pedipalps.
Class Arachnida

l Class Arachnida includes spiders,

scorpions, mites, and ticks.

50 µm
Class Arachnida

l Two tagmata:
l Cephalothorax
l Chelicerae
l Pedipalps
l 4 pairs walking legs
l Abdomen
Class Arachnida - Order Araneae

l Most spiders – order

Araneae – have 8 simple
eyes that can detect light and
motion.
l Some hunting & jumping
spiders may form images.
l Sensory setae detect air
currents, web vibrations, and
other stimuli.
l Spider’s vision usually poor,
so awareness of environment
depends largely on cuticular
mechanoreceptors such as
sensory setae.
Class Arachnida - Order Araneae

l All are predaceous,

mostly on insects.
l Many spin a web used
for prey capture.
l Some chase & catch
prey.
l Injected venom
liquefies and digests
the tissues which is
sucked into spider’s
stomach.
Class Arachnida - Order Araneae
l Two or three pairs of
spinnerets contain
microscopic tubes that
run to silk glands.
l Liquid scleroprotein
secretion hardens as it is
extruded from spinnerets.
l Silk threads are very
strong and will stretch
considerably before
breaking.
l Silk is used for orb webs,
lining burrows, forming
egg sacs, and wrapping
prey.
Class Arachnida - Order Araneae
l Breathe by book lungs and/or tracheae.
l Book lungs unique to spiders - parallel air pockets extend into
blood-filled chamber.
l Air enters chamber through a slit in body wall.
l Tracheae system is less extensive than in insects.
l Transports air directly to tissues.
l Tracheal systems of arthropods represent a case of evolutionary
convergence.
Class Arachnida - Order Araneae

l In spiders and insects, Malpighian tubules

serve as excretory structures.
l Potassium, other solutes, and waste are secreted
into tubules.
l Rectal glands reabsorb the potassium and water,
leaving wastes and uric acid for excretion.
l Conserves water and allows the organisms to live
in dry environments.
l Many spiders have coxal glands, modified
nephridia, at the base of legs.
Class Arachnida - Order Araneae

l Reproduction - before mating, male

stores sperm in pedipalps.
l Mating involves inserting pedipalps into the
female genital openings.
l A courtship ritual is often required before the
female will allow mating.
l Eggs may develop in a cocoon in the web or
may be carried by female.
l Young hatch in about two weeks and may
molt before leaving the egg cocoon.
Class Arachnida - Order Araneae
l Are spiders really dangerous?
l Spiders are allies of humans in our battle with insects.
l American tarantulas rarely bite, and bite is not dangerous.
l Species of black widow spiders are dangerous.
l Venom is neurotoxic.
l Brown recluse spider has hemolytic venom that destroys tissue
around the bite.
l Some Australian and South American spiders are the most
dangerous and aggressive.
Class Arachnida - Order Scorpiones

l Scorpions – order
Scorpiones – feed on
insects & spiders which
they seize with their
pedipalps.
l The last segment
contains a bulbous base
and a curved barb that
injects venom.
l Scorpions are viviparous
or ovoviviparous –
females brood young
within their reproductive
tract.
Class Arachnida - Order Opiliones

l Harvestmen – order
Opiliones – differ from
spiders in that the
abdomen and
cephalothorax are
broadly joined rather
than constricted.
l Only two eyes
l Abdomen shows
segmentation
l Long legs end in tiny
claws.
Class Arachnida - Order Acari

l Mites and ticks – order Acari – have a fused

cephalothorax & abdomen.
l Mites are tiny – 1mm or less.
l Some feed on plant juices and can be major pests.
l Several species of ticks carry diseases such as
Lyme disease.
Class Arachnida - Order Acari

l House dust mites - free-

living and often cause
allergies.
l Spider mites - one of many
important agricultural pest
mites that suck out plant
nutrients.
Class Arachnida - Order Acari

l Hair follicle mite

Demodex - harmless
but other species
cause mange in
domestic animals.
l Human itch mite
causes intense
itching.
Class Arachnida - Order Acari

l Tick species of
Ixodes carry Lyme
disease.
l Tick species of
Dermacentor
transmit Rocky
Mountain spotted
fever.
l Cattle tick transmits
Texas cattle fever.
Subphylum Myriapoda
l Subphylum
Myriapoda includes
these classes:
l Chilopoda
(centipedes)
l Diplopoda
(millipedes)
l Pauropoda
(pauropods)
l Symphyla
(symphylans)
l Use trachea to
transport air.
l Excretion usually by
Malpighian tubules.
Class Chilopoda

l Centipedes – class
chilopoda – contain
a few or many
segments each
(except the first
behind the head and
the last two) with a
pair of jointed legs.
l Last pair of legs has
a sensory function.
Class Chilopoda

l Head appendages:
l One pair antennae
l One pair
mandibles
l One or two pairs
of maxillae.
l Dorsoventrally
flattened.
Class Chilopoda

l Sexes are separate.

l Some lay eggs (oviparous).
l Some have live young (viviparous).

l Young like little adults – no metamorphosis.

Class Chilopoda

l Centipedes live in moist environments.

l They are carnivores, feeding on insects &
worms.
l Prey is killed with poison claws on the first
segment.
Class Diplopoda

l Millipedes (Class
Diplopoda) have
two pairs of legs on
each segment.
l Head has one pair
each of antennae,
mandibles, &
maxillae.
l Body is more
cylindrical.
Class Diplopoda

l Millipedes live in dark, moist places – under

rocks or logs.
l Most are herbivores, feeding on decayed plant
matter or occasionally living plants.
l Slow moving, coil up when disturbed.
l Toxic or repellent fluids secreted when
disturbed.
Class Diplopoda

l After copulation,
female lays eggs in a
nest and guards
them.
l Larvae have only
one pair of legs per
segment.
Class Pauropoda

l Live in moist soil, leaf litter, decaying

vegetation, or under bark and debris.
l Least well known of myriapods.
l Soft-bodied, small (2 mm or less).
l Approximately 500 species.
l Head lacks true eyes, has branched
antennae, and a pair of sense organs.
Class Pauropoda

l 12 trunks segments bear

9 pairs of legs but none
on the first or last 2
segments.
l One tergal plate covers
two segments.
l Lack tracheae, spiracles,
and circulatory system.
l Probably most closely
related to diplopods.
Class Symphyla
l Live in humus, leaf mold, and
debris.
l Male Scutigerella places a
spermatophore at end of a
stalk.
l Female stores the sperm in
special pouches.
l Removes and smears
eggs with sperm before
attaching them to moss or
lichen.
l Young hatch with only 6 or 7
pairs of legs.
Class Symphyla
l Small (2–10 mm) with centipede-like
bodies.
l Soft-bodied with 14 segments - 12
segments bear legs and one bears a
pair of spinnerets.
l Antennae are long and unbranched.
l About 160 species are known.
l Eyeless with sensory pits at base of
antennae.
l Tracheal system connects to a pair
of spiracles on the head and tracheal
tubes to the anterior only.
Phylogeny

l Relationships between subphyla are debated.

l Taxon of Pancrustacea, which includes
hexapods and crustaceans, is well-supported.
l Phylogenies using molecular data rarely
support grouping Myriapoda with
Pancrustacea.
l There is support for placement of Myriapoda
as the sister taxon for Cheliceratae.
Phylogeny
l Biologists assume that the ancestral arthropod
had a segmented body with one pair of legs
per segment.
l Evolution caused adjacent segments to fuse
and to make body regions.
l Hox gene studies indicate that the first five
segments fused to form the head tagma in all
four extant subphyla.
l In spiders, Hox gene studies indicate that the
entire prosoma corresponds to the head of
other arthropods.
Phylogeny

l Genetic studies have been helpful in

understanding the evolution of uniramous and
biramous appendages.
l Molecular evidence repeatedly places hexapods
with crustaceans even though hexapods have
uniramous appendages and crustaceans have
biramous appendages.
l Leads to the question: Did uniramous
appendage development evolve more than
once?
Adaptive Diversification
l In contrast to annelids, arthropods have
pronounced tagmatization by fusion of somites.
l Those with primitive characters have
appendages on each somite.
l Derived forms are specialized.
l Modification of exoskeleton and appendages
allowed variation in feeding and movement.
l Adaptations made possible by cuticular
exoskeleton and small size fostered high
diversity.
Phylogeny
l Subphylum Trilobita
l Subphylum Chelicerata
l Class Merostomata
l Class Pycnogonida
l Class Arachnida
l Subphylum Myriapoda
l Class Diplopoda
l Class Chilopoda
l Class Pauropoda
l Class Symphyla
l Subphylum Crustacea
l Subphylum Hexapoda