22/08/2020

SYSTEMA
ARTHROPODA
SYSTEMATICS LECTURE
Lecture Handout Chapter 19 – Trilobites, Chelicerates,
and Myriapods
(Selected slides from PPT of Chapter 20 – Crustaceans
Hickman et al. 15ed) Chapter 21 - Hexapods

Characteristics
“Arthropodization”
• Soft cuticle of the arthropod ancestors was stiffened
by deposition of protein and chitin
• Joints had to provide flexibility
• Sequence of molts was necessary to allow for growth
• Molting required hormonal control
• Hydrostatic skeleton function was lost
– Coelom regressed and was replaced by open sinuses
• Motile cilia were lost
Characteristics
Phylum Arthropoda
• Contains over 3/4 of all known species
• Approximately 1,100,000 species of arthropods have
been recorded
• Rich fossil history dating to late Precambrian
• Eucoelomate protostomes with well-developed
organ systems
• Segmented
• Molecular analyses indicate annelids and arthropods
evolved from different ancestors

19-3 19-4

Characteristics
• Sizes range from the Japanese crab (four meters in
leg span) to the 0.1 mm long follicle mite
• Abundance and wide ecological distribution makes
them the most diverse animal group
• Some are agents of disease and compete with
humans for food
• Others are beneficial
• All modes of feeding occur among arthropods but
most are herbivorous
Characteristics
Relationships among arthropod subgroups
• Divided into subphyla based on relationships between
subgroups
• Groupings among subphyla based on molecular data
• Centipedes, millipedes, pauropods, and symphylans are
placed in subphylum Myriapoda
• Insects are placed in subphylum Hexapoda
• Spiders, ticks, horseshoe crabs and their relatives form
subphylum Chelicerata
• Lobsters, crabs, barnacles, and others form subphylum
Crustacea

19-5 19-6

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19-7 19-8

Characteristics Great Diversity and Abundance of Arthropods

• Also included in Crustacea are tongue worms • Versatile Exoskeleton

• Extinct trilobites are placed in subphylum Trilobita
• Relationships are controversial
• “Mandibulate hypothesis”
– Myriapods, hexapods, and crustaceans more closely related
due to a shared mouthpart, the mandible
• Molecular evidence of a close relationship between
hexapods and crustaceans unites subphylum
Crustacea with subphylum Hexapoda in clade
Pancrustacea
– Cuticle is highly protective and jointed, providing mobility
– Consists of inner thick procuticle and outer thin epicuticle
– Procuticle has an exocuticle secreted before a molt and an
endocuticle secreted after molting
– Both layers of procuticle contain chitin bound with protein
– Procuticle is lightweight, flexible, and protects against
dehydration
– Chitin content varies from 40% of the procuticle in insects to
as much as 80% in crustaceans

19-9 19-10

Great Diversity and Abundance of Arthropods

– Calcium salts responsible for hardness of procuticle of

lobsters and crabs
– Cuticle is laminated and further hardened by tanning
– Cuticle is thin between segments, allowing for movement
at the joints
– Ecdysis, or molting
• Process of shedding outer covering and growing a new, larger one
– Arthropods typically molt four to seven times
• Weight is a limit to ultimate body size

19-11 19-12

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Great Diversity and Abundance of Arthropods Great Diversity and Abundance of Arthropods

• Segmentation and Appendages for Efficient • Air Piped Directly to Cells

Locomotion
– Primitive pattern
• Linear series of similar somites with jointed appendages
– Many somites may be fused or combined into specialized
groups called tagmata
• Highly Developed Sensory Organs
– Appendages often highly specialized for division of labor
– Limb segments are hollow levers with internal striated
muscles
– Appendages may function in sensing, food handling,
walking, or swimming
– Terrestrial arthropods
• Use an efficient tracheal system for oxygen transport
– Aquatic arthropods
• Respire via various forms of gills
– Eyes vary from simple light sensitive ocelli to a compound
mosaic eye
– Other sensory structures for touch, smell, hearing,
balancing, and chemical reception

19-13 19-14

Great Diversity and Abundance of Arthropods Subphylum Trilobita

• Complex Behavior Patterns
– Arthropods surpass most other invertebrates in complex
and organized activities
– Most behavior is innate but some is learned
• Use of Diverse Resources through Metamorphosis
– Many arthropods undergo metamorphic changes leading
to different larval and adult stages
– Larvae and adults feed on different organisms and occupy
different habitats
• Avoid competition
History of an Ancient Group
• Trilobites arose before the Cambrian, flourished, and
then became extinct 200 million years ago
• Trilobed body shape due to a pair of longitudinal
grooves
• Bottom dwellers and probably were scavengers
• Ranging from 2 to 67 centimeters long
• Could roll up like pill bugs
• Exoskeleton contained chitin strengthened by calcium
carbonate

19-15 19-16

Subphylum Trilobita
• Body was divided into a cephalon, trunk, and
pygidium
– Cephalon was a fusion of segments
– Trunk varied in number of somites
– Pygidium was fused into a plate
• Cephalon bore antennae, compound eyes, a mouth,
and jointed appendages
• Each body somite except the last had a pair of
biramous appendages
• One of the branches of biramous appendage was
fringed
A B – May have been a gill
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19-17 19-18

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Subphylum Chelicerata
Characteristics
• Chelicerates have six pairs of cephalothoracic
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19-20

Subphylum Chelicerata
Class Merostomata: Subclass Eurypterida
• Eurypterids (giant water scorpions) were the largest of all
fossil arthropods
– 3 m in length
• Fossils date from Ordovician to Permian periods
• Resemble both marine horseshoe crabs and terrestrial
scorpions
• Head: 6 fused segments, 6 pairs of appendages, simple
and compound eyes, chelicerae, pedipalps, and 4 pairs of
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walking legs
• Abdomen: 12 segments and spike-like telson

19-21 19-22

Subphylum Chelicerata Subphylum Chelicerata

Class Merostomata: Subclass Xiphosurida, Horseshoe • Walk with walking legs and swim with abdominal
Crabs plates
• Modern horseshoe crab nearly unchanged from • Feed at night on worms and small molluscs
ancestors in the Triassic period
• 5 species in 3 genera survive • During mating season, come to shore at high tide to
• Most live in shallow water mate
• Structures • Females burrow into sand to lay eggs
– Unsegmented carapace covers body in front of a broad
abdomen and a telson • Males follow to add sperm before eggs are covered
– Cephalothorax has 5 pairs of walking legs and a pair of • Young larvae hatch and return to sea at next high tide
chelicerae
– Abdomen bears six pairs of broad, thin, appendages fused in • Larvae are segmented and resemble trilobites
the median line
– Book gills exposed on some abdominal appendages
– Carapace has 2 compound and 2 simple eyes
19-23 19-24

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Subphylum Chelicerata
Class Pycnogonida: Sea Spiders
• All have small, thin bodies
• Approximately 1,000 species
• Some species duplicate somites, and may have five
or six pairs of legs
• Males may have a pair of legs (ovigers) to carry
developing eggs
• Many have chelicerae and palps
• Mouth, at the tip of a proboscis, sucks juices from
cnidarians and soft-bodied animals

19-25 19-26

I
Subphylum Chelicerata Subphylum Chelicerata
• Most have four simple eyes Class Arachnida
• Simple dorsal heart • Great diversity
• No excretory and respiratory systems • Most are free living and more common in warm, dry
• Digestive system sends branches to the legs regions
• Most gonads are in the legs Structures
• Occupy all oceans but most common in polar waters • Divided into two tagmata: a cephalothorax and an
• Some suggest that pycnogonids belonged to an abdomen
early-diverging arthropod lineage • Cephalothorax bears a pair of chelicerae, a pair of
pedipalps, and 4 pairs of walking legs
• No antenna and mandibles

-
19-27 19-28
I, I

Subphylum Chelicerata Subphylum Chelicerata

• Most are predaceous and have claws, fangs, poison
glands, or stingers
• Sucking mouthparts ingest fluids and soft tissues
from bodies of their prey
• Spiders have spinning glands
• A few spiders may have a segmented abdomen, a
primitive character
• Pedipalps of males are modified, sometimes
elaborately, for sperm transfer
• Over 80,000 species have been described
• Scorpions appeared on land in the Silurian, mites and
spiders by the end of the Paleozoic Era
• Most harmless to humans and provide essential
control of injurious insects
• Some spiders are venomous and can cause pain or
death in humans
• Ticks may carry human diseases
• Mites can be crop pests

19-29 19-30

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Subphylum Chelicerata
Order Araneae: Spiders Opisthosoma Prosoma
(abdomen) (cephalothorax)
• Approximately 40,000 species
• Body consists of an unsegmented cephalothorax and
abdomen joined by a slender pedicel
• Anterior appendages are a pair of chelicerae with
terminal fangs
• Pair of pedipalps have sensory functions and are used
by males to transfer sperm
• Basal parts of pedipalps used to handle food Chelicerae Pedipalp
• Four pairs of walking legs terminate in claws
• All are predaceous, mostly on insects

19-31 19-32

Subphylum Chelicerata
• Injected venom liquefies and digests the tissues which is
sucked into spider’s stomach
• Breathe by book lungs and/or tracheae
• Book lungs unique to spiders
• Parallel air pockets extend into blood-filled chamber
• Air enters chamber through a slit in body wall
• Tracheae system is less extensive than in insects
– Transports air directly to tissues
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19-33 19-34

Subphylum Chelicerata Subphylum Chelicerata

• In spiders and insects, Malpighian tubules serve as
excretory structures
• Potassium, other solutes, and waste are secreted
into tubules
• Rectal glands reabsorb the potassium and water,
leaving wastes and uric acid for excretion
• Conserves water and allows the organisms to live in
dry environments
• Many spiders have coxal glands, modified nephridia,
at the base of legs
• Sensory Systems
– Most spiders have eight simple eyes, each with a
lens, optic rods, and a retina
• Detect movement and may form images
– Sensory setae detect air currents, web vibrations,
and other stimuli
– Spider’s vision usually poor
• Awareness of environment depends largely on
cuticular mechanoreceptors such as sensory
setae

19-35 19-36

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Subphylum Chelicerata
• Web-Spinning Habits
– Spinning silk critical ability for spiders and some other
arachnids
– Two or three pairs of spinnerets contain microscopic tubes
that run to silk glands
– Liquid scleroprotein secretion hardens as it is extruded
from spinnerets
– Silk threads are very strong and will stretch considerably
before breaking
– Silk is used for orb webs, lining burrows, forming egg sacs,
and wrapping prey

19-37 19-38

Subphylum Chelicerata
• Reproduction
– Before mating, male stores sperm in pedipalps
– Mating involves inserting pedipalps into the female genital
openings
– A courtship ritual is often required before the female will
allow mating
– Eggs may develop in a cocoon in the web or may be carried
by female
– Young hatch in about two weeks and may molt before
leaving the egg cocoon

19-39 19-40

Subphylum Chelicerata
• Are spiders really dangerous?
– Most people fear spiders without good reason
– Spiders are allies of humans in our battle with insects
– American tarantulas rarely bite, and bite is not dangerous
– Species of black widow spiders are dangerous
• Venom is neurotoxic
– Brown recluse spider
• Hemolytic venom that destroys tissue around the bite
– Some Australian and South American spiders are the most
dangerous and aggressive

19-41 19-42

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19-43
19-45
Subphylum Chelicerata
Order Solpugida: Sun or Camel Spiders
• Sun, wind or camel spiders
• Nonvenomous, shred their prey with large
chelicerae
• Range from 1 cm to 15 cm
• Common in America, Middle East, Asia, Africa
22/08/2020
Subphylum Chelicerata
Order Scorpionida: Scorpions
• More common in tropical and subtropical zones but
occur in temperate areas
• Approximately 1,400 species worldwide
• Nocturnal and feed largely on insects and spiders
• Sand-dwellers locate prey by detecting surface waves
with leg sensillae
• Appendages attached to cephalothorax
– Pair of medial eyes and 2–5 lateral eyes
• Preabdomen has 7 segments
• Postabdomen has long, slender tail of five segments
that ends in a stinging apparatus
19-44
Subphylum Chelicerata
• Under the abdomen are comblike pectines
– Explore the ground and aid in sex recognition
• Stinger on last segment has venom that varies from
mildly painful to dangerous
• Ovoviviparous or viviparous and produce from 6 to
90 young
• Perform complex mating dances
– In some species the male stings the female on
pedipalp or on edge of cephalothorax
19-46
Subphylum Chelicerata
Order Opiliones: Harvestmen
• Harvestmen or daddy longlegs
• Common, particularly in tropical regions
• Approximately 5,000 species worldwide
• Unlike spiders, abdomen and cephalothorax join
broadly without a narrow pedicel
• Can lose most of their eight long legs without ill
effect
• Chelicerae are pincerlike
• Mostly scavengers
19-48
8

Subphylum Chelicerata
Order Acari: Ticks and Mites
• Medically and economically the most important
arachnids
• About 30,000 species have been described
• Both aquatic and terrestrial
– Inhabit deserts, polar areas, and hot springs
• Most mites are less than 1 millimeter long
• Ticks may range up to 2 cm
• Complete fusion of cephalothorax and abdomen
• No sign of external segmentation
19-49
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19-51
19-53
22/08/2020
Subphylum Chelicerata
• Mouthparts on tip of the anterior capitulum
• Chelicerae on each side help pierce, tear, or grip food
• Other mouthparts include pedipalps with a fused
base, hypostome, rostrum, and tectum
• Adult mites and ticks possess 4 pairs of legs
• Transfer sperm directly or by spermatophores
• Egg hatches, releasing a six-legged larva
– Eight-legged nymphal stages follow
19-50
Subphylum Chelicerata
• House dust mites
– Free-living and often cause allergies
• Spider mites
– One of many important agricultural pest mites that suck
out plant nutrients
• Chiggers
– Larval Trombicula mites
– Feed on dermal tissues and cause skin irritation
• Hair follicle mite Demodex
– Harmless but other species cause mange in domestic
19-52
19-54
9
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Subphylum Chelicerata
• Human itch mite
– Causes intense itching
• Tick species of Ixodes
– Carry Lyme disease
• Tick species of Dermacentor
– Transmit Rocky Mountain spotted fever
• Cattle tick
– Transmits Texas cattle fever

19-55 19-56

Subphylum Myriapoda
Characteristics
• Myriopods include
– Chilopoda (centipedes)
– Diplopoda (millipedes)
– Pauropoda (pauropods)
– Symphyla (symphylans)
• Use trachea to transport
• Excretion usually by Malpighian tubules

19-57 19-58

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Subphylum Myriapoda

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Class Chilopoda

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• Centipedes
• Found under logs, bark and stones
• Carnivorous, eating earthworms, cockroaches, and

- other insects
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Yenomte,igs – Common in bathrooms and damp cellars
• Most harmless to humans
– Few large, tropical centipedes are dangerous
• Approximately 3,000 species worldwide
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19-59 19-60

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Subphylum Myriapoda
Characteristics
E.. F"nti.,p.. &llaDNI~ • Terrestrial and have flattened bodies with up to 177
segments
• Each segment except the one behind the head and last
two
– Bears a pair of jointed legs, the last pair of which
serves a sensory function
• Appendages of first body segment form poison claws
• Head has one pair of antennae, a pair of mandibles,
and one or two pairs of maxillae
• Eyes on either side of the head consist of groups of
ocelli
19-61 19-62

Subphylum Myriapoda Subphylum Myriapoda

• Salivary glands empty into anterior end of straight Reproduction
digestive tract
• Two pairs of Malpighian tubules empty into the hind • Sexes separate with unpaired gonads and paired
intestine ducts
• Elongated heart has pair of arteries in each somite • Some lay eggs and others are viviparous
– Ostia provide return flow of hemolymph
• Young resemble adults and do not undergo
• Pair of spiracles in each somite allows air to diffuse
through branched air tubes of the tracheae metamorphosis
• Arthropod nervous system includes a portion that
serves as a visceral nervous system

19-63 19-64

Subphylum Myriapoda
Class Diplopoda
Natural History
• Millipedes
• Less active than centipedes
– Walk with graceful rather than wriggling motion
• Most eat decayed plants but a few eat living plant
tissue
• Most are slow moving and roll into a coil for defense
• Some secrete toxic or repellant fluids from special
repugnatorial glands on side of body
• More than 10,000 species of worldwide
19-65 19-66

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Subphylum Myriapoda Subphylum Myriapoda

Characteristics
• Cylindrical bodies with 25 to more than 100 segments
• Short thorax consists of 4 segments, each bearing one
pair of legs
• Head has 2 clusters of simple eyes and a pair each of
antennae, mandibles, and maxillae
• Each abdominal somite has 2 pairs of spiracles
opening into air chambers and tracheal air tubes
• Two genital apertures located toward anterior end
Reproduction
• Appendages of seventh segment specialized as
copulatory organs
• After copulation, female lays eggs in a nest and
guards them
• Larvae have only one pair of legs per segment

19-67 19-68

Subphylum Myriapoda Subphylum Myriapoda

Class Pauropoda
• 12 trunks segments bear 9 pairs of legs but none on
Life History the last 2 segments
• Live in moist soil, leaf litter, decaying vegetation, or – A tergal plate covers each of the two segments
under bark and debris • Lack tracheae, spiracles, and circulatory system
• Least well known of myriapods • Probably most closely related to diplopods
Characteristics
• Soft-bodied, small (2 mm or less)
• Approximately 500 species
• Head lacks true eyes, has branched antennae, and a
pair of sense organs

19-69 19-70

Subphylum Myriapoda
Class Symphyla
Life History and Reproduction
• Live in humus, leaf mold, and debris
• Male Scutigerella places a spermatophore at end of a
stalk
• Female stores the sperm in special pouches
– Removes and smears eggs with sperm before attaching
them to moss or lichen
A • Young hatch with only 6 or 7 pairs of legs
19-71 19-72

12

Subphylum Myriapoda
Characteristics
• Small (2–10 mm) with centipede-like bodies
• Soft-bodied with 14 segments
– 12 segments bear legs and one bears a pair of spinnerets
• Antennae are long and unbranched
• About 160 species are known
• Eyeless with sensory pits at base of antennae
• Tracheal system connects to a pair of spiracles on the
head and tracheal tubes to the anterior only
19-73
Phylogeny and Adaptive Diversification
• Biologists assume that the ancestral arthropod had a
segmented body with one pair of legs per segment
• Evolution caused adjacent segments to fuse and to
make body regions
• Hox gene studies indicate that the first five segments
fused to form the head tagma in all four extant
subphyla
• In spiders, Hox gene studies indicate that the entire
prosoma corresponds to the head of other
arthropods
19-75
Phylogeny and Adaptive Diversification
– Leads to the question: Did uniramous appendage
development evolve more than once?
– Numbers of appendages per segment is another variable
character among arthropods that lends itself to more
testing
19-77
22/08/2020
Phylogeny and Adaptive Diversification
Phylogeny
• Relationships between subphyla are debated
• Taxon of Pancrustacea, which includes hexapods and
crustaceans, is well-supported
• Phylogenies using molecular data rarely support
grouping Myriapoda with Pancrustacea
• There is support for placement of Myriapoda as the
sister taxon for Cheliceratae
19-74
Phylogeny and Adaptive Diversification
• Sea spiders remain within subphylum Chelicerata
because Hox gene studies have found that their head
appendage arose from the region of the head that
corresponds to the second segment
• Genetic studies have been helpful in understanding
the evolution of uniramous and biramous
appendages
– Molecular evidence repeatedly places hexapods with
crustaceans even though hexapods have uniramous
appendages and crustaceans have biramous appendages.
19-76
Phylogeny and Adaptive Diversification
Adaptive Diversification
• In contrast to annelids, arthropods have pronounced
tagmatization by fusion of somites
• Those with primitive characters have appendages on
each somite
– Derived forms are specialized
• Modification of exoskeleton and appendages allowed
variation in feeding and movement
• Adaptations made possible by cuticular exoskeleton
and small size fostered high diversity
19-78
13
 22/08/2020

Phylogeny and Adaptive Diversification Crustaceans

Classification
• Subphylum Trilobita • Over 67,000 living species
• Subphylum Chelicerata • Insects and crustacea compose over 80% of all
– Class Merostomata named animal species
– Class Pycnogonida • Probably most abundant animals in the world are
– Class Arachnida members of the copepod genus Calanus
• Subphylum Myriapoda • Divided into 5 classes
– Class Diplopoda – Current molecular phylogenies do not support the
– Class Chilopoda monophyly of all classes
– Class Pauropoda • Former members of phylum Pentastomida placed in
– Class Symphyla class Maxillopoda, subclass Pentastomida
• Subphylum Crustacea
• Subphylum Hexapoda
19-79 20-80

Overview
- -
j _ --TL • Pentastomids
– Also called tongue worms
:"r: ~:, – Parasites of vertebrates, living in lungs or nasal cavities
– Closely related to fish lice

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20-81 20-82

Subphylum Crustacea Subphylum Crustacea

General Nature of a Crustacean • Primitive crustaceans may have up to 60 segments;
• Main distinguishing characteristic of crustaceans derived crustaceans have fewer
– Two pairs of antennae
• Tagmata are usually head, thorax, and abdomen
• Head also has a pair of mandibles and 2 pairs of
maxillae – Not homologous across taxa
• One pair of appendages on each of the additional • In most one or more thoracic segments are fused with
segments the head as a cephalothorax
– Some segments may lack appendages • Arrangement of tagmata in Malacostraca is the ancestral
– All appendages, except perhaps first antennae, are biramous
plan
– Head has 5 fused somites
– Thorax has 8
– Abdomen has 6
20-83 20-84

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Subphylum Crustacea
Cephalolhorax Abdomen
3 segments 6 segments • Anterior end is a non-segmented rostrum
• Telson, with the last abdominal somite and uropods,
forms a tail
• Dorsal covering is the carapace
– May cover most of body or just cephalothorax

Telson
Uropod

Walking ~s Swimmerets
20-85 20-86

Subphylum Crustacea
Form and Function
• External Features
– Secreted cuticle is made of chitin, protein, and calcareous
material
– Heavy plates have more calcareous deposits
• Joints are soft and thin, allowing flexibility
– Dorsal tergum and ventral sternum are plates on each
somite lacking a carapace
– Telson is not a somite
• Bears anus and may be homologous to the pygidium
– In some species, telson may bear a pair of processes, the
caudal furca
– Gonopores may be at base of appendages, at the tail, or on
somites without legs
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20-87 20-88

Subphylum Crustacea
GUI
• Appendages
– Members of Malacostraca and Remipedia have appendages
on each somite
– Other classes may not bear appendages on abdominal
Co,a}- Protopod
somites Basis

– Specialization of appendages based on the basic biramous

plan
– Maxilliped has a basal protopod, a lateral exopod, and a Endopod
medial endopod
– Endites are medial processes, exites are lateral processes
and epipod is an exite on the protopod
– Appendages represent serial homology
• Have evolved a wide variety of walking legs, mouthparts,
swimmerets, etc. from modification of the basic biramous
appendage
20-89 20-90

15
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20-91 20-92
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Subphylum Crustacea Subphylum Crustacea

• Internal Features
– Muscular and nervous systems and segmentation exhibit
metamerism of annelid-like ancestors
– Hemocoel
• Persistent blastocoel that becomes filled with blood
• Coelomic compartments remain as end sacs of
excretory organs and gonads
• Coelomates only in technical sense of the term
• Muscular System
– Striated muscles
• Make up a major portion of crustacean body
– Most muscles arranged as antagonistic groups
• Flexors draw a limb toward the body and extensors
straighten a limb out
– Abdominal flexors of a crayfish allow it to swim backward
– Strong muscles located on each side of stomach control
the mandibles

20-93 20-94

Subphylum Crustacea
• Respiratory System
– Smaller crustaceans may exchange gases across thinner
areas of cuticle
– Larger crustaceans use featherlike gills for gas exchange
– Decapod carapace overlaps the gill cavity, leaving anterior
and posterior openings
– “Bailer” of 2nd maxilla draws water over gill filaments
........... .......... O,.-Ow9bl ..... ~ . . . , , , , _ ....... – Gills may project from pleural wall, the articulation of
OllflCI ~ ...... ....._.....
thoracic legs, or thoracic coxae

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Subphylum Crustacea
• Circulatory
– Open circulatory system
• No system of veins to separate blood from interstitial fluid
• Hemolymph exits heart through arteries
• Passes to hemocoel to return to the heart via sinuses
– Dorsal heart
• Single-chambered sac of striated muscle
– Valves in the arteries prevent backflow of hemolymph
– Hemolymph conducted to gills, if present, for oxygen and
carbon dioxide exchange
– Hemolymph may be colorless, reddish, or bluish
• Contains ameboid cells that may help prevent clotting
– Hemocyanin and/or hemoglobin are respiratory pigments

20-97 20-98

Subphylum Crustacea
• Excretory System Bladder

– Antennal or maxillary glands

Labyrinth
– Called green glands in decapods
– End sac of antennal gland has a small vesicle and a spongy
labyrinth
– Labyrinth connects by an excretory tubule to dorsal
bladder that opens to exterior pore
– Hydrostatic pressure within a hemocoel provides a force
for filtration of fluid into the end sac
– Resorption of salts and amino acids occurs as the filtrate
passes the excretory tubule and bladder
• Mainly regulates the ionic and osmotic composition of
body fluids

20-99 20-100

Subphylum Crustacea Subphylum Crustacea

• Nervous and Sensory Systems
– Nitrogenous wastes excreted across thin areas of cuticle in
the gills
– Freshwater crustaceans constantly threatened by over-
dilution with water
• Gills must actively absorb Na+ and Cl-
– Marine crustaceans have urine that is isosmotic with blood
20-101
– More fused ganglia than in other arthropods
– Pair of supra-esophageal ganglia connects to eyes and two
pairs of antennae
– Neuron connectives join this brain to the subesophageal
ganglion
• Supplies nerves to mouth, appendages, esophagus, and antennal
glands
– Double ventral nerve cord has a pair of ganglia for each somite
to control appendages
– Eyes and statocysts are largest sensory organs
– Tactile hairs occur on the body, especially on chelae,
mouthparts and telson
– Chemical sensing of taste and smell occurs in hairs on
antennae and mouth
20-102

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Subphylum Crustacea
– Statocyst opens at base of first antenna in crayfish
– Statocyst lined with sensory hairs that detect position of
grains of sand
– Compound eyes, made of many units called ommatidia
– Cornea focuses light down the columnar ommatidium
– Distal retinal, proximal retinal, and reflecting pigment cells
form a sleeve around each ommatidium
– Each ommatidium detects a restricted area of objects, a
mosaic, in bright light
– In dim light, the distal and proximal pigments separate and
produce a continuous image

Day-adapted Night-adapted
20-103 20-104

Subphylum Crustacea Subphylum Crustacea

• Reproduction, Life Cycles, and Endocrine Function
– Diversity of Reproduction
• Barnacles are monoecious but generally cross-fertilize
• In some ostracods, males are scarce and reproduction
is by parthenogenesis
• Most crustaceans brood eggs in brood chambers, in
brood sacs attached to the abdomen, or attached to
abdominal appendages
• Crayfishes develop directly without a larval form
• Most crustaceans have a larva unlike the adult in form,
and undergo metamorphosis
• The nauplius is a common larval form with uniramous
first antennae, and biramous second antennae and
mandibles that all aid in swimming
– Appendages and somites are added in a series of
molts
• Metamorphosis of a barnacle proceeds from a free-
swimming nauplius to a larva with a bivalve carapace
and finally to a sessile adult with plates

20-105 20-106

0
Egg
Subphylum Crustacea

* Nauplius
• Ecdysis
– Necessary for a crustacean to increase in size the
• Exoskeleton does not grow
– Physiology of molting affects reproduction, behavior, and
many metabolic processes
– Underlying epidermis secretes cuticle
– Outermost epicuticle is made of a very thin lipid-
impregnated protein
Protozoea
– Most of the cuticle is composed of several layers of the
procuticle
– Exocuticle, beneath the epicuticle, contains protein,
calcium salts, and chitin

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Subphylum Crustacea
– Endocuticle has a heavily calcified principal layer and an
uncalcified membranous layer
– Molting animals grow in the intermolt phases, or instars
– Soft tissue increases in size until there is no space within
the cuticle
– When body fills the cuticle, animal is in the premolt phase
– Epidermal cells enlarge before ecdysis

20-109 20-110

Subphylum Crustacea
– Secrete a new epicuticle and then begin secreting a new
exocuticle
– Enzymes released into the area above new epicuticle
dissolve the old endocuticle
– When only the old exocuticle and epicuticle remain, the
animal swallows water to expand and burst the old cuticle
– Soft new cuticle stretches and then hardens with the
deposition of inorganic salts
– Molting occurs often in young animals and may cease in
adults

20-111 20-112

Subphylum Crustacea Subphylum Crustacea

• Hormonal Control of Ecdysis
– Temperature, day length, or other stimuli trigger central
nervous system to begin ecdysis
– Central nervous system decreases production of molt-
inhibiting hormone by the X-organ
– Promotes release of molting hormone from the Y-organs
which promotes ecdysis
• Other Endocrine Functions
– Removing eyestalks accelerates molting and prevents color
changes to match background
– Hormones from neurosecretory cells in eyestalk control
dispersal of cell pigment
– Neurosecretions from pericardial organs stimulate increase
in heartbeat
– Androgenic glands in male amphipods stimulate
expression of male characteristics

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Subphylum Crustacea
• Feeding Habits Dorsal tooth Pyloric stomach
– Same fundamental mouthparts are adapted to a wide
array of feeding habits Cardiac ...,,, - - - Dorsal cecum
– Suspension feeders generate water currents in order to stomach -,,.1."";l!!!!!i>-- lntestine
feed on plankton, detritus ,and bacteria
Ventral cecum
– Predators consume larvae, worms, crustaceans, snails, and Gastrolith
fishes
– Scavengers eat dead animal and plant matter
– Crayfishes have a two-part stomach
• Gastric mill grinds up food in 1st compartment

20-115 20-116

Brief Survey of Crustaceans

Class Remipedia
• Only 10 described species
• All found in caves connected to the sea
• Primitive features include 25–38 segments with
similar, paired, biramous, swimming appendages
• Antennules also biramous
• Maxillae and maxillipeds are prehensile and
specialized for feeding
• Swimming legs are directed laterally rather than
ventrally as is found in copepods and cephalocarids

20-117 20-118

Brief Survey of Crustaceans Brief Survey of Crustaceans

Class Cephalocarida
• Only 9 species described
• Live in coastal bottom sediments from intertidal
zones to 300 meters depth
• Thoracic limbs and 2nd maxillae are very similar
• Lack eyes, a carapace, and abdominal appendages
• True hermaphrodites and unique in discharging eggs
and sperm through same duct
Class Branchiopoda
• Over 10,000 species, 4 orders
– Order Anostraca, includes fairy shrimp and brine
shrimp
• Lack a carapace
– Order Notostraca, includes tadpole shrimp
• Carapace forms a large dorsal shield
– Order Conchostraca, includes clam shrimp
• Enclosed by a bivalved carapace
– Order Cladocera, includes water fleas
• Carapace encloses the body but not the head

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Brief Survey of Crustaceans

• In all flattened, leaf-like legs serve as respiratory
organs, assist in suspension feeding, and, (except for
cladocerans), locomotion
• Most are freshwater
• Important component of freshwater zooplankton
• May reproduce by parthenogenesis to rapidly boost
summer populations and then by sexual
reproduction with the onset of unfavorable
conditions
A Tadpole shrimp
(order Notostraca) C Daphnia
(order Diplostraca,
suborder Cladocera)
20-121 20-122

Brief Survey of Crustaceans Brief Survey of Crustaceans

• Fertilized eggs highly resistant to cold
– Critical for winter survival of population
• Cladocerans: Development mostly direct
• Other branchiopods: Gradual metamorphosis
Class Ostracoda
• 6,000 known species
• Most are dioecious
• Enclosed in a bivalve carapace
• 0.25–8.0 mm long
• Fusion of trunk somites
– Thoracic appendages are reduced to two or one
• Most are benthic or climb onto plants, but some are
planktonic, parasitic, or burrowing
• Widespread in both marine and freshwater habitats
• Development by gradual metamorphosis

20-123 20-124

Brief Survey of Crustaceans

Class Maxillopoda
• General body plan
– Five cephalic, six thoracic and four abdominal somites plus
a telson
• No appendages on abdomen
• When present, the eye is unique in structure and
called a maxillopodan eye

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Brief Survey of Crustaceans Brief Survey of Crustaceans

Subclass Mystacocarida
• Less than 0.5 mm long
• Live in interstitial water between sand grains
• 10 species have been described from around the
world
• Primitive in several characteristics
Subclass Copepoda
• Third in numbers of species
• Lack a carapace and retain simple, median, nauplius
eye in the adult
• Single pair of uniramous maxillipeds and four pairs of
flattened, biramous, thoracic swimming appendages
• A major articulation separates the posterior from the
anterior, appendage-bearing portion of the body
• Antennules often longer than other appendages

20-127 20-128

Brief Survey of Crustaceans Brief Survey of Crustaceans

• Parasitic forms highly modified and reduced • Development is indirect
– Often unrecognizable as arthropods • Some highly modified parasites have unusual
• Free-living copepods may be the dominant consumer metamorphoses
• Marine copepod Calanus is most abundant organism
in zooplankton by biomass
• Cyclops and Diaptomus important elements of
freshwater plankton
• Some free-living copepods are intermediate hosts of
human parasitic tapeworms and nematodes

20-129 20-130

Brief Survey of Crustaceans

Subclass Tantulocarida
• Only recently described
• Approximately 12 species
• Tiny copepod-like ectoparasites of deep-sea benthic
crustaceans
• No head appendages beyond one pair of antennae in
sexual females
• Likely alternate between a parthenogenetic cycle and
a bisexual cycle with fertilization
• Tantalus larvae penetrate cuticle of host by mouth
tube
• Abdomen and all thoracic limbs are lost during
metamorphosis to an adult
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Brief Survey of Crustaceans Brief Survey of Crustaceans

Subclass Branchiura
Subclass Pentastomida
• Lack gills
• Tongue worms
• Most are ectoparasites of marine and freshwater fish • Comprise about 130 species of parasites of
• 5–10 mm long vertebrate respiratory systems
• Broad, shield-like carapace, compound eyes, four • Most infect reptile lungs, a few infect air sacs of birds
biramous thoracic swimming appendages, and a short or mammals
unsegmented abdomen • More common in tropical regions extending out to
• Second maxillae are modified as suction cups to North America, Europe, and Australia
attach to host fish • Range from 1 to 13 cm in length
• Development is direct • Chitinous cuticle regularly molted

20-133 20-134

Brief Survey of Crustaceans

• Five protuberances on the anterior end provide the
phylum name
– Four of the protuberances bear claws
– Fifth contains the mouth and two pairs of hooks
• Simple straight digestive system
• Nervous system has paired ganglia along the ventral
nerve cord
• Lack any circulatory, excretory, or respiratory organs
• Sexes are separate

20-135 20-136

Brief Survey of Crustaceans Brief Survey of Crustaceans

• Females larger than males
– Produces millions of eggs that pass up the host trachea, are Subclass Cirripedia
swallowed and leave host with feces • Includes barnacles in order Thoracica and three orders
• Larvae hatch out as oval, tailed creatures with four of burrowing or parasitic forms
stumpy legs • Adults are sessile and attach directly (acorn barnacles)
• Most life cycles require an intermediate vertebrate or by a stalk (goose barnacles)
host, usually a fish or reptile • Carapace surrounds body and secretes a set of
• After ingestion by an intermediate host calcareous plates
– Larva penetrates intestine and migrates until it changes to a • Head is reduced, abdomen is absent, and thoracic
nymph legs are long with hairlike setae
– Nymph becomes encapsulated and remains dormant until
eaten • Jointed cirri bear setae and extend from the plates to
• When eaten, juvenile migrates to lung, feeds on blood feed on small particles
and tissue, and matures • In barnacles in intertidal zones, plates close to protect
against dessication

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Brief Survey of Crustaceans

• Most non-parasitic barnacles are hermaphroditic and
undergo metamorphosis during development
– Most hatch as nauplii and become cyprid larvae with a
bivalve carapace and compound eyes
– Attach to substrate by first antennae and adhesive glands
– Secrete calcareous plates, lose eyes, and transformation of
swimming appendages to filtering cirri
– Parasitic forms may have a kentrogon stage that injects
cells into the hemocoel of host

20-139 20-140

Brief Survey of Crustaceans

Class Malacostraca
• Largest and most diverse class of Crustacea with over
20,000 species
• Contains three subclasses, 14 orders, and many
suborders

Order Isopoda
• Only truly terrestrial crustaceans
• Also have marine and freshwater forms
• Dorsoventrally flattened, lack a carapace, and have
sessile compound eyes
20-141 20-142

Brief Survey of Crustaceans

• 1st pair of thoracic limbs are maxillipeds
• Remaining thoracic limbs lack exopods
• Abdominal appendages bear gills, except for uropods
• Common land forms include sow bugs and pill bugs
• Cuticle lacks protection of insect cuticle
– Must live in moist habitats
• Some isopods are highly modified as parasites of fishes or
crustaceans
• Development typically direct but may be metamorphic in
parasitic forms

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Brief Survey of Crustaceans

Order Amphipoda
• Amphipods resemble isopods
– Lack a carapace, have sessile compound eyes, and one pair
of maxillipeds
• However, they are compressed laterally, and gills are
in the thoracic region
• Abdominal and thoracic limbs are grouped for
jumping and swimming
• Many are marine, others are beach-dwelling,
freshwater, or parasitic
• Development is direct

20-145 20-146

20-147 20-148

Brief Survey of Crustaceans

Order Euphausiacea
• Approximately 90 species
• Includes important ocean plankton called krill
• Carapace does not completely enclose gills
• Lack maxillipeds and all limbs have exopods
• Most are bioluminescent with a light-producing organ
called a photophore
• Form a major component of the diet of baleen whales
and of many fishes
• Eggs hatch as nauplii
• Development is direct

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Brief Survey of Crustaceans

Order Decapoda
• 5 pairs of walking legs and 3 pairs of maxillipeds
• In crabs, first pair of walking legs form pincers
• Range from a few millimeters to the Japanese crab with a
4 m leg-span
• Approximately 18,000 species
• Includes crayfishes, lobsters, crabs, and true shrimp
• Crabs have a broader cephalothorax and reduced
abdomen, compared to crayfish or lobsters

20-151 20-152

Phylogeny and Adaptive Diversification Phylogeny and Adaptive Diversification

Phylogeny
• Remipedia appear to be the most primitive of
Crustacea
• Fossils of an arthropod in the Mississippian are likely
the sister group to remipedians
• One theory is that each modern somite represents two
ancestral somites that fused together, forming the
biramous appendage
• The pentastomids were placed in Ecdysozoa because
of the similarities in larvae, molting of the cuticle, and
sperm morphology
• Phylogenies based on rRNA genes and affirmed in
sequences of mitochondrial DNA indicate that
pentastomids are crustaceans
20-153
Adaptive Diversification
• Crustaceans are unquestionably the dominant
arthropod in marine environments
• They also share dominance in freshwater
environments with the insects
• The class Malacostraca is most diverse and members
of Copepoda are most abundant
• Copepods are particularly successful as parasites of
both vertebrates and invertebrates
20-154

Phylogeny and Adaptive Diversification Phylogeny and Adaptive Diversification

Classification • Class Ostracoda
– Subclass Maxillopoda
• Class Remipedia – Subclass Mystacocarida
• Class Cephalocarida – Subclass Copepoda
• Class Branchiopoda
– Order Anostraca – Subclass Tantulocarida
– Order Notostraca – Subclass Branchiura
– Order Cladocera – Subclass Pentastomida
– Order Conchostraca – Subclass Cirripedia

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Phylogeny and Adaptive Diversification

• Class Malacostraca
– Order Isopoda
– Order Amphipoda
– Order Euphausiacea
– Order Decapoda

20-157 21-158

Diversity and Characteristics

Subphylum Hexapoda
• Members named for the presence of six legs
– All legs are uniramous
• Have 3 tagmata
– Head
– Thorax
– Abdomen
• Appendages attach to head and thorax
• Two classes within Hexapoda
– Entognatha
– Insecta
·-
--
..........,___, ---
-•--
21-159 21-160

Diversity and Characteristics Diversity and Characteristics

• Characteristics of Entognatha
– Small group characterized by bases of mouthparts • Characteristics of Insecta
enclosed within the head capsule
– Enormous class whose members have ectognathous
– 3 orders: Protura, Diplura, and Collembola mouthparts, however, bases of mouthparts lie outside the
• Protura and Diplura head capsule
– Tiny, eyeless, and inhabit soils or dark, damp places
– Pterogotes: winged insects
• Collembola
– Commonly called springtails because of ability to leap – Apterogotes: wingless insects
– Animal 4 mm long may leap 20 times its body length
– Members live in soil, decaying plant matter, on freshwater
pond surfaces, and along seashore
– Can be very abundant, reaching millions per hectare

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Class Insecta Class Insecta

• Diversity
– Most diverse and abundant of all arthropods
– Number of known species estimated at 1.1 million, but
estimated that there may be as many as 30 million species
worldwide
– Continued evolution among modern insects
• Fossil record indicates they are a stable group
– Play major medical and economic roles with humans, and
also play critical ecological roles
• Characteristics
– Ectognathous mouthparts and often two pair of wings on
the thoracic region of the body
– Range from less than 1 mm to 20 cm in length
• Larger insects are tropical
• Distribution
– Found in nearly all habitats except the sea
– Common in freshwater, brackish water, and salt marshes
– Abundant in soils, forest canopies, and can be found in
deserts and wastelands
– Most animals and plants have insects as parasites externally
and internally
• Adaptive Traits
– Flight and small size makes insects widely distributed
– Well-protected eggs withstand rigorous conditions and are
readily dispersed
– Wide variety of structural and behavioral adaptations gains
them access to every possible niche

21-163 21-164

Class Insecta Class Insecta

• Adaptability
– Most structural modifications are in wings, legs, antennae,
mouthparts, and alimentary canal
– Hard, protective exoskeleton well-adapted to life in desert
regions
– Exoskeleton holds in water, a factor that allows desert
survival
External Form and Function
• Exoskeleton composed of complex plates, or sclerites,
connected by hinge joints
– Muscles attaching sclerites allow precise movement
• Rigidity is due to scleroproteins and not mineral matter
– Allows for lighter weight body necessary for flight
• More homogenous in tagmatization than the variable
crustaceans
• Cuticle of a somite is composed of a dorsal notum, a
ventral sternum and a pair of lateral pleura

21-165 21-166

Class Insecta
• Head
– Usually equipped with pair of large compound
eyes
– One pair of antennae varies greatly in
• Function in touch, taste, hearing
– Mouthparts consist
• Labrum
• Pair of mandibles and maxillae,
• Labium
• Hypopharynx

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Class Insecta
Silkwormlarvll
• Thorax
– Consists of the prothorax, mesothorax and metathorax
– Each section has a pair of legs
• Wings

-- – If two pairs present: located on the mesothorax and

metathorax
– Consist of a double membrane
– Veins serve to strengthen the wing
• Vein pattern used to identify insect taxa

-- 21-169 21-170

Class Insecta
• Legs
– Walking legs end in terminal pads and claws
– Hindlegs of grasshoppers and crickets are enlarged for
jumping
– Mole crickets have front legs adapted for burrowing in
ground
– Forelegs of praying mantis allow it to grasp prey
– Honeybees have leg adaptations for collecting pollen

21-171 21-172

Femur Trochant8f Coxa

Hlndleg
(lateral view)

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Class Insecta
• Abdomen
– 9 to 11 segments
– Last is reduced to a pair of cerci
– Larval and nymphal forms may have abdominal appendages
lacking in adults
– External genitalia usually at end of abdomen
• Variations in Body Form
– Land beetles are thick and shielded
– Aquatic beetles are streamlined
– Cockroaches are flat and live in crevices
– Antennae vary widely from long to short, plumed to
knobbed
21-175 21-176

Class Insecta
• Locomotion: Walking
– Insects walk using first and last leg on one side and middle
leg on the opposite side in alteration with the reverse
• Provides stability
– A water strider has non-wetting footpads that do not break
the surface water tension
• Power of Flight
– Insect wings not homologous with bird and flying mammal
wings
– Insect wings are outgrowths of cuticle from the
mesothoracic and metathoracic segments
– Recent fossil evidence suggests insects may have evolved
fully functional wings over 400 million years ago
21-177 21-178

Class Insecta
– Recent fossil evidence suggests insects may have evolved
fully functional wings over 400 million years ago
– Most flying insects have 2 pairs of wings
• Diptera (true flies) have one pair
– Halteres are reduced wings that provide the fly with
balance during flight
– Non-reproductive ants and termites are wingless
– Lice and fleas have also lost wings
• Modifications of Wings
– Wings for flight are thin and membranous
– The thick and horny front wings of beetles are protective
– Butterflies have wings covered with scales
– Caddisflies have wings covered with hairs

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Class Insecta
• Flight Muscles of Insects
– Direct flight muscles attach to wing directly
– Indirect flight muscles alter the shape of the thorax to
cause wing movement
– Wing is hinged on a pleural process that forms a fulcrum
– Insects cause the upstroke with indirect muscles that pull
the tergum downward
– Dragonflies and cockroaches contract direct muscles to
pull the wing downward
– Bees, wasps and flies arch the tergum to cause the
downstroke indirectly
– Beetles and grasshoppers use a combination of direct and
indirect muscles to move wings

21-181 21-182

Class Insecta
• Flight Muscle Contraction
– Synchronous muscle control uses a single volley of nerve
impulses to stimulate a wing stroke
– Asynchronous muscles stretch antagonistic muscle and

ilJr
cause wing to contract in response
• Require occasional nervous stimulation
– Potential energy can be stored in resilient tissues
OQcl and l'ICMCI llghl rni.--. hlirtalligl'Crnuldll – Wing beats may vary
• 4/second in butterflies
ollooAlsMd~ ol'-Mdmodga9

• 1000/second in midges
C

21-183 21-184

Class Insecta Class Insecta

• Wing Thrust
Internal Form and Function
– Direct flight muscles alter the angle of wings to twist
• Digestive System
leading edge to provide thrust – Foregut
– Figure-8 movement moves insect forward • Mouth with salivary glands, esophagus, crop, and gizzard
• Some digestion, but no absorption, occurs in crop as
– Fast flight requires long, narrow wings and a strong tilt, as salivary enzymes mix with food
in dragonflies and horse flies • Gizzard grinds food before it enters the midgut
– Midgut
• Primary site of digestion and absorption
• Ceca may increase digestive and absorptive area
– Hindgut
• Primarily a site for water absorption

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Class Insecta
– Most feed on plant tissues or juices and are herbivorous or
phytophagous
– Many caterpillars are specialized to eat only certain species
of plants
– Some ants and termites cultivate fungus gardens for food
– Many beetles and other insect larvae eat dead animals and
are saprophagous
– Some species are predaceous on other insects or other
animals

21-187 21-188

Class Insecta
– Many species are parasitic as adults and/or larvae
– Many parasitic insects, in turn, have parasites, a condition
called hyperparasitism
– Parasitoids live inside a host and eventually kill the host
• Important in pest control

21-189 21-190

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Class Insecta
• Mouthparts
– Sucking mouthparts form a tube to pierce tissues of
animals or plants
– Houseflies and blowflies have sponging mouthparts
• Soft lobes at the tip absorb food
– Biting mouthparts can seize and crush food

A B
AClewanl!Ptwxnan, .. , B J H ~ P Y n s

21-193 21-194

Class Insecta
• Circulation
– Tubular heart in the pericardial cavity moves hemolymph
forward through dorsal aorta
– Heartbeat is a peristaltic wave
– Accessory pulsatile organs help move hemolymph into
wings and legs
– Hemolymph composed of plasma and amebocytes but
does not function in oxygen transport in most insects
– In some insects, particularly aquatic immature in low
oxygen environments
• Hemoglobin is present and functions in oxygen transport

21-195 21-196

Class Insecta
• Gas Exchange
– Terrestrial animals are faced with the dilemma of
exchanging gases while preventing water loss
– Tracheal system
·-
• Network of thin-walled tubes that branch throughout the insect
body
• Evolved independently of that of other arthropod groups
– Spiracles open to the tracheal trunks
• 2 spiracles on thorax and 7–8 on abdomen
– Valve on the spiracle
• Reduces on water loss and may serve as a dust filter

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Class Insecta Class Insecta

– Tracheae are composed of a single layer of cells lined with
cuticle that is shed at each molt
– Spiral thickenings of cuticle, called taenidia, prevent the
tracheae from collapsing
– Tracheae branch out into fluid-filled tubules called
tracheoles that reach individual body cells
– System provides gas transport without use of oxygen-
carrying pigments
– Diving beetles use abdominal hairs to maintain a bubble
under wings, an “artificial gill”
– Mosquito larvae use short breathing tubes to snorkel
surface air
– Air sacs in insects are dilated tracheae without taenidia
– Contraction of muscles in the jaw or limbs causes increased
pressure inside the exoskeleton
• Elevated pressure causes contraction of tracheae for
exhalation
– Muscular movements may assist in moving air in and out of
air sacs
– Very small insects
• Transport gases by simple diffusion
– Aquatic insect nymphs
• May use tracheal gills or rectal gills

21-199 21-200

Class Insecta
• Excretion and Water Balance
– Insects and spiders utilize Malpighian tubules in conjunction
with rectal glands
– Malpighian tubules vary in number but join between the
midgut and hindgut
– Blind ends of tubules float freely in hemocoel bathed in
hemolymph
– Potassium is actively secreted into the tubules ..... H,O ;ii: KHCO,. H,O
• Other solutes follow gradient
– Main waste product is uric acid '°"""'' ~ .
• Flows across at upper end that is mildly alkaline
– In the lower end of the tubule
• Potassium combines with CO2 and is reabsorbed
..., ....
– Rectal glands reabsorb chloride, sodium and water
• Wastes pass out of body ?/
21-201 21-202

Class Insecta Class Insecta

• Nervous System • Sense Organs
– Resemble that of larger crustaceans, with fusion of ganglia – Many insects have keen sensory perception
– Most sense organs are microscopic and located in body
– Some have a giant fiber system wall
– A stomadeal system corresponds to the autonomic system – Different organs respond to mechanical, auditory,
of vertebrates chemical, visual, and other stimuli
– Neurosecretory cells in brain function to control molting – Mechanoreception
and metamorphosis • Touch, pressure, vibration, etc. are detected by sensilla
– May be a single hair-like seta or a complex organ
– Distributed widely over antennae, legs, and body

21-203 21-204

34
 22/08/2020

Class Insecta Class Insecta

– Auditory Reception
• Sensitive setae (hair sensilla) or tympanal organs detect
airborne sounds
• Tympanal organs occur in Orthoptera, Hemiptera and
Lepidoptera
• Organs in legs can detect vibrations of substrate
– Chemoreception
• Usually bundles of sensory cell processes located in
sensory pits
• May occur on mouthparts, antennae, and legs
• Some insects can detect odors several kilometers away
• Feeding, mating, habitat selection, and host-parasite
relationships are mediated through chemical senses

21-205 21-206

Class Insecta
– Visual Reception
• 2 types of eyes: simple and compound
• Honeybee studies indicate that ocelli monitor light
intensity but do not form images
• Compound eyes may contain thousands of ommatidia
Clmmaldum

-
--
-"
er,,,aone""""

.........
– Ommatidia structure similar to that of crustaceans
• Insects can see simultaneously in almost all directions
– Image is myopic and fuzzy
COmpound'!"

21-207 21-208

Class Insecta Class Insecta

• Flying insects have a higher flicker-fusion rate – Other Senses
– Distinguish 200–300 flashes per second • Insects are very sensitive to temperature, especially
• A bee can distinguish ultraviolet light but cannot detect cells in antennae and legs
shades of red • Insects also detect humidity, proprioception, gravity,
and other physical properties

21-209 21-210

35
 22/08/2020

Class Insecta Class Insecta

• Neuromuscular Coordination • Reproduction
– Active insects require excellent neuromuscular – Parthenogenesis occurs predominantly in some Hemiptera
coordination and Hymenoptera
– Arthropod muscles are cross-striated – Sexual reproduction is the norm
– Sexes are separate
– Strength of muscle is related to its cross-sectional area
– Sexual Attraction
– A flea can jump 100 times its length by storing energy in an
• Female moths secrete a pheromone to attract males
elastic resilin protein from a great distance
• Fireflies use flashes of light to detect mates
• Some insects use sounds, color signals, and other
courtship behaviors

21-211 21-212

Class Insecta
– Fertilization usually internal
– Sperm may be released directly or packaged into
spermatophores
• Spermatophores are result of an evolutionary transition from marine
to terrestrial existence
• May be transferred both without copulation and during copulation.
– Female may only mate once and store sperm to fertilize eggs
throughout her life
– Females may lay a few eggs and provide care of young, or lay
huge numbers
– Butterflies and moths must lay eggs on the host plant if the
caterpillars are to survive
– Wasps may have to locate a specific species that is the only
host to their young
21-213 21-214

~ ~
\Ad~;:butterlly
mago)

Egg (ow)

Caterpillar
(lan,a)
Buuertty
emerging
from chrysalis
OAcCle't~PnolDSffilct

21-215 21-216

36
 22/08/2020

Class Insecta
• Metamorphosis and Growth
– Various forms of metamorphosis produce degrees of
change among different insect groups
– Most insects change form after hatching from egg
– Each stage between molts is called an instar
– Insects develop wings during the last stage

A
OOr--lc.ww:.8

21-217 21-218

.
.,

21-219 21-220

Class Insecta Class Insecta

– Ametabolous (Direct) Development
• Silverfish and springtails have young similar to adults
except in size and sexual maturation
• Stages are egg-juveniles-adult
• Wingless insects
– Hemimetabolous Metamorphosis
• Some insects undergo a gradual metamorphosis
• Grasshoppers, cicadas, mantids, true bugs, mayflies and
dragonflies exhibit this metamorphosis
• Young are called nymphs
• Bud-like growths in early instars show where the adult
wings will eventually develop
• Stages are egg-nymph-adult

21-221 21-222

37

Class Insecta
– Holometabolous Metamorphosis
• About 88% of insects undergo complete metamorphosis
• Separates the physiology of larval growth, pupal
differentiation, and adult reproduction
• Larvae and adults often live in completely different
environments
– No competition
• After several larval instars
– Larval moth or butterfly becomes a pupa inside a cocoon or
chrysalis
– Pupae often pass the winter in this stage
– Final molt occurs and the adult emerges in spring
• Stages are egg-larva-pupa-adult
21-223
Class Insecta
• Corpora allata produces juvenile hormone
• Molting continues as long as juvenile hormone
(neotenine) is sufficiently present
• In later instars, the corpora allata releases less juvenile
hormone
• When juvenile hormone reaches a low level, the larva
molts to become a pupa
• Cessation of juvenile hormone production in the pupa
leads to an adult at the last molt
• In hemimetabolous insects, cessation of juvenile
hormone occurs in the last nymphal instar
• In adults, the corpora allata becomes active again in
normal egg production
• The prothoracic glands degenerate in adult insects and
adults do not molt
21-225
Class Insecta
– Defense
• Protective coloration, warning coloration, and mimicry
are protective adaptations
• Stink bugs and others have repulsive odors and tastes
• Some insects are aggressive (e.g., bees and ants)
• The monarch caterpillar incorporates a poisonous
substance from its food plant, milkweed
• The bombardier beetle can spray an attacking enemy
with irritating chemicals
21-227
22/08/2020
Class Insecta
– Physiology of Metamorphosis
• Hormones regulate insect metamorphosis
• Brain and nerve cord ganglia produce brain hormone or
ecdysiotropin
• Neurosecretory cells send axons to the corpora cardiaca
that stores and ultimately releases these hormones
• Brain hormone circulates in the hemolymph to the
prothoracic gland in the head or prothorax
• Prothoracic gland produces molting hormone or
ecdysone in response
• Ecdysone starts the molting process
21-224
Class Insecta
– Diapause
• Period of dormancy in the annual life cycle that is
independent of conditions
– Winter dormancy: hibernation
– Summer dormancy: estivation
• Any stage (eggs, larvae, pupae, or adults) may remain
dormant to survive adverse conditions
• Internally controlled but may be triggered by
environmental cues such as day length
• Always occurs at end of an active growth stage
– Insect is then ready for another molt
• Many larvae do not develop beyond this point until
spring in spite of mild temperatures
21-226
21-228
38
 22/08/2020

Class Insecta
• Behavior and Communication
– Insects respond to many environmental stimuli
– Responses are governed by both physiological state of the
animal and its nerve pathways
– Many behaviors are complex sequences of responses
– Most behavior is innate but some involve simple learning

21-229 21-230

Class Insecta
– Pheromones
• Chemicals secreted by one individual to affect the
behavior of another individual
• Attract the opposite sex, trigger aggregation, fend off
aggression, and mark trails
• Bees, wasps, and ants can recognize nestmates and
signal an alarm if strangers enter the nest
• Can be used to trap insects to monitor populations

21-231 21-232

Class Insecta
– Sound Production and Reception
• Sounds are used as warning devices, advertisement of
territory, and courtship songs.
• Crickets chirp for courtship and aggression
• Male cicada vibrates paired membranes on abdomen to
attract females

– Tactile Communication
• Involves tapping, stroking, grasping, and antennae
touching
• Some beetles, flies and springtails use bioluminescence
• Some female fireflies mimic another species’ flash pattern
to attract males and then eat them

21-233 21-234

39
 22/08/2020

Class Insecta Class Insecta

Honeybees
– Social Behavior • Have a few male drones, a fertile female queen and many
• Some social communities are temporary and female workers
uncoordinated • Drones develop by parthenogenesis
• Others are highly organized and depend on chemical • Development of a fertile queen requires ingestion of “royal
and tactile communication jelly”
• Caste differentiation is common in most organized • Queen secretes “queen substance” to prevent workers
social groups from maturing or feeding larvae royal jelly
• A honeybee hive may contain 60,000–70,000 individuals
• Scouts inform workers on location of food

21-235 21-236

Class Insecta
Termites
• Fertile king and queen fly away to start a new colony
– Mate and lose wings
• Immature members are wingless and become workers
and soldiers
• Soldiers have large heads and defend colony
• Reproductive individuals secrete inhibiting pheromones
that produce sterile workers
• Nymphs feed from each other in trophallaxis, thus
spreading the pheromone about

21-237 21-238

Class Insecta
• Worker castes also produce worker and soldier
substances
• Drops in these pheromone levels result in more of the
needed caste developing in the next generation

A
A . O J H ~ J .IIODrJamnL~

21-239 21-240

40
 22/08/2020

Class Insecta
Ants
• Differ from termites
– Ants are darker, hard-bodied, and have thread-like
waist
• In ant colonies, the male ant dies after mating
• Ants have wingless soldiers and workers, and often
have variations of these castes
• Ants have also evolved striking patterns of “economic”
behavior: making slaves, fungus farming, sewing nests
together, tool use, and herding

21-241 21-242

Insects and Human Welfare Insects and Human Welfare

• Beneficial Insects
– Insects produce honey, beeswax, silk and shellac – Dead animals rapidly consumed by fly maggots
– Of more economic importance, bees pollinate $10 – Insects are critical components of most food
billion worth of food crops in the U.S. annually chains and an important food source for many fish
– Pollinating insects and flowering plants are tightly and birds
co-evolved
– Predaceous and parasitoid insects are vital in
controlling many pest insect populations

21-243 21-244

Insects and Human Welfare

• Harmful Insects
– Harmful insects eat and destroy plants and fruits
– Nearly every cultivated crop has several insect
pests
• Requires substantial money for insect control
– Bark beetles, spruce budworms, the gypsy moth
and others are serious forest pests.
– Insects also destroy food, clothing, and property

21-245 21-246

41
 22/08/2020

Insects and Human Welfare

– Medically important insects include vectors for
disease agents
– 10% of all arthropod species are parasites or
“micropredators”
– Warble and bot flies attack humans and domestic
livestock
– Malaria is carried by Anopheles mosquitos
• Most common major world disease
– Yellow fever and lymphatic filariasis are also
mosquito-borne

21-247 21-248

Insects and Human Welfare

– Fleas carry plague, a disease that changed human
history in the Middle Ages
– Lice carry typhus fever
– The tsetse fly carries African sleeping sickness
– The newest viral plague to hit North America, the
West Nile virus, is carried by mosquitos

00rJameSLC$lff

21-249 21-250

Insects and Human Welfare Insects and Human Welfare

• Control of Insects
– Broad-spectrum insecticides damage beneficial
insect populations along with targeted pest
– Some chemical pesticides persist in the
environment and accumulate as they move up the
food chain
– Some strains of insects have evolved a resistance
to common insecticides
– Biological control
• Use of natural agents, including diseases, to suppress
an insect population
– Bacillus thuringiensis
• Bacterium that controls lepidopteran pests
• Gene coding for the “B.t.” toxin has been introduced to
other bacteria and transferred to crop plants
– Some viruses and fungi may be economical
pesticides
– Natural predators or parasites of insect pests can
be raised and released to control pest
– Release of sterile males can eradicate the few
insect species that only mate once

21-251 21-252

42

Insects and Human Welfare
– Pheromones can monitor pests and hormones
may play a role in disrupting the life cycle
– Integrated pest management
• Combined use of all possible, practical techniques listed
above, to reduce reliance on chemical insecticides
21-253
Phylogeny and Adaptive
Diversification
– Some phylogenies support a sister-taxon
relationship between them, but others indicate
that hexapods arose within Crustacea
– Future studies may show that subphylum Crustacea
is paraphyletic
– Phylogenies that support hexapods arising from
within Crustacea, find that hexapods are most
similar to brachiopod, cephaplocarid, and
remipedian crustaceans
21-255
Phylogeny and Adaptive
Diversification
– Within Hexapoda, Entognatha is the sister taxon
to class Insecta
– However, some research indicates that
entognathous mouthparts may have evolved
several times and that some entognathans are
closer to insects than to other entognathans
21-257
22/08/2020
Phylogeny and Adaptive Diversification
• Phylogeny
– Understanding of the relationships among
arthropods has changed over the past decade
– Using molecular data, members of former
subphylum Uniramia are now divided between
subphylum Myriapoda and Hexapoda
– The nature of the relationship, however, between
hexapods and crustaceans is not well understood
21-254
Abdominal --;;;..---...;:..
limbs
A B
21-256
Phylogeny and Adaptive
Diversification
• Adaptive Diversification
– The first terrestrial arthropods were scorpions and
millipedes that appeared in the Silurian period
– The ancestral insect had a head and trunk of
similar somites
– Insects have specialized the first three post-
cephalic somites as thorax and lost the remaining
appendages
– Some modern apterygote orders have abdominal
styli that are considered vestigial legs
21-258
43

Phylogeny and Adaptive
Phylogeny and Adaptive Diversification
Diversification
– Recent fossil evidence suggests winged insects
were in existence about 400 million years ago
– Ancestral flying insects may have derived from
aquatic insects or insects with aquatic juveniles;
the external gills on their thorax may be the
derivative of wings
– Metamorphosis also distinguished insects by the
Permian period
21-259
22/08/2020
– Hemimetabolous metamorphosis, chewing
mouthparts and cerci group the Orthoptera,
Dermaptera, Isoptera and Embioptera
– Hemimetabolous metamorphosis and sucking
mouthparts group the Thysanoptera, Hemiptera,
Homoptera and perhaps the Psocoptera, Zoraptera,
Mallophaga and Anoplura
– Other orders have holometabolous metamorphosis
and are the most specialized
21-260
44