Zoology 111 – Invertebrate Zoology Jules Raposas

Origin of Life
• Review: Creationism
-universe and all life forms created by a human being
• Theory of Spontaneous Generation -life generates -
itself from non-living matter -soil, mud →
microbes trapped in curve
salamanders, frogs, worms, etc.
-life arises from non-living material, in addition to
Microscopy
biogenesis
-enabled natural scientists to examine beyond what
• Plagues in History they see from the naked eye
• Robert Hooke → cellulae
*science debunks myths
→dynamic • Anton van Leeuwenhoek → animalcules -middle
ages in Europe used molded bread in healing
• Debunked by Francesco Redi (1668) -maggots wounds
spontaneously arise from spoiled meat • Alexander Fleming
-discovered Staphylococcus by accident

• John Needham tried to debunk this (1745)

-Penicillium notatum
→penicillin
-no knowledge of -mass produced for the wounded in World War II
extremophiles (strict anaerobes) -up to date, used for many infections

• Lazzaro Spallanzani also made an experiment in 1765 *antibiotic resistance is a problem nowadays
-“antibac” cause mutations to microorganisms
→triclosan
*conjugation
-possess genetic changes from consuming bacteria
→plasmid

-E. coli – produces vitamin K for blood clotting to

prevent diarrhea
• Louis Pasteur
-swan neck experiment

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Abiogenesis -Oparin-Haldane Hypothesis

 -primitive atmosphere was reducing → -genes are activated when the organism is in a group
synthesis of organic compounds -ex: bioluminescence
• Cellularization
Typical Prokaryotic Cell
• Group behavior genes Classification Schemes
-bacteria -archaea extremophiles
o Monera (bacteria)
o Protista (invertebrates)
o Plants
o Animals
o Fungi
• 6-Kingdom Model
-plasmid is
o Eubacteria
o Archaebacteria
independent in encoding and expressing genes
o Protista
→endosymbiosis o Plants
-bacteriophages o Animals
o Fungi
Prokaryotes to Eukaryotes
• 8-Kingdom Model
• Lynn Margulis – wife of Carl Sagan -
endosymbiosis
-large number of major lineages without isolated
*blue green algae = cyanobacteria
representatives
→ancestor of plants
-bacteria comprise majority of life’s current -
-photosynthetic eukaryotes and archaea are closely related -entire
-invaginations and enfoldings continent of life forms

Invertebrates
-group of animals lacking a backbone -do not develop a
notochord during embryogenesis

Importance
• Ecological
o Waste recycling
o Ecosystem structure
Unicellular to Multicellular o Pollination
• Colony formation o Food webs
-organism is multicellular by colonial stage -each o Bioindicators
protist individual could remain together after division • Agricultural
-ex: colony of Volvox sp. o Pest control
o Parasites
• Scientific
• 5-Kingdom Model
o Baseline data on biodiversity

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o Model species for researches

o Robotics
• Medical
o Human parasites
 o Medical researches
o Source of biomedical substances
• Commercial
o Commercially important compounds
o Aesthetic purposes
▪ Tubipora musica

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Principles of Systematics -discovery, description, interpretation of diversity

-science of organismal diversity
-study of biological diversity and its evolutionary history System
-scheme/method of procedure or classification to put -represented by phylogenetic tree
order (for orderliness and organization) *animal diversity -relationship of one organism to another
– one of the greatest manifestations
-invertebrates are not a homogenous group of phyla, an *phylogenetic tree
assortment, falling into a number of phyla -same as family trees
*invertebrates account for 99% for all species of animals i. Common ancestor
ii. Which organism branched
Goals of Systematic Zoologists out from a common ancestor
• To discover all species of animals (living or extinct) iii. Which organisms are closely
• To reconstruct their evolutionary relationships related to each other
• To classify them accordingly *Darwin recognized the major source of
evidence in homology
Classifying Organisms *Ontogeny – history of the development of
Theme an organism through its entire life
-diversity may be incomprehensible but there are themes -Ernst Haeckel
1. Bauplan -recapitulation/biogenetic law:
-basic design/defining characteristics ontogeny repeats phylogeny
-ex: arthropods, segmented body, exoskeleton, 3. Functional Principles
jointed appendages, lice, crabs, and grasshoppers a. Convergent: evolved due to adaptation to
2. Evolutionary Relationships similar environment
a. Theory of Common Descent -ex: flight, swimming
-all plants and animals came from the same b. Divergent: one ancestral stock, different traits
ancestor -evolution toward different traits in closely
-debunked by horizontal gene transfer, related species
conjugation
Cell Number
• Acellular
-do not resemble anything that resembles a
multicellular embryo
-gray area
-some organisms have parts that are considered to be
non-living because they are not made up of cells
-plasmids, viruses
b. Phylogeny • Unicellular
-origin and diversification of organisms • Multicellular
-evolutionary history -collectively known as metazoa

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Symmetry nervous and sensory tissues and organs at one end of

-regular arrangement of body structure relative to body the animal, resulting in distinct posterior and anterior
axis end
-general body form 2. Radial Symmetry
1. Bilateral Symmetry -2 reasonably equal halves by any cut that pass
-left and right through the center; no front or back end 3. Asymmetrical
-approximate mirror images of each other -highly -no body axis or no plane of symmetry 4.
correlated with cephalization: concentration of Metamerism or Segmentation
 -serial repetition of body segments along →blastomeres (morula)
longitudinal axis of the body -ends with formation of blastula
5. Tagmatization • Radial Cleavage
-fusion or combination of metameres into • Spiral Cleavage
specialized functional units -cleavage planes are always oblique to the
polar axis of the embryo
Developmental Pattern
-growth of an organism
1. Germ Layer
-stem cells
-give rise to different tissues and/or organ systems
• Diploblastic (endoderm and ectoderm)
• Triploblastic (endoderm, mesoderm, and
ectoderm)
2. Type of Coelom
-fluid-filled cavity lying between the gut and the
outer body wall musculature and lined with tissue
derived from embryonic mesoderm
• Acoelomate – flatworms → lack an internal
body 5. Fate
• Eucoelomate – annelids (segmented) → true
coelom
• Pseudocoelomate – roundworms → region
between outer body wall and endoderm of of the Blastomere
the gut is a fluid-filled cavity • Regulative/Intermediate Cleavage -
deuterostomes
3. Mechanism of Coelom Formation • Schizocoely –
• Mosaic/Determinate Cleavage
gradual enlargement of a split in mesoderm
(protostomes) -development stops when 1 cell is excised
-protostomes
• Enterocoely – invagination of archenteron into
6. Fate of the Blastopore
the primitive blastocoel (dueterostomes)
-opening of archenteron
4. Cleavage Pattern
→division of cells in the early development -rapid • Deuteroustomes → anus
cell cycles with no significant growth • Protostomes → mouth

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 -relationship
-symbiont: benefits
a. Ectosymbiont – live near
b. Endosymbiont – live within the body c.
Mutualism
d. Commensalism
e. Parasitism

Systematics
1. Taxonomy/Nomenclature
2. Classification
a. Artificial – morphology, anatomy, other
aspects
b. Natural – morphoanatomy, have assumptions
Lifestyle and Habitat on progression of evolution, inferences on
1. Ecological Preference relationships
a. Freshwater – lower salt concentration and 3. Evolutionary Relationship
other dissolved solids
b. Terrestrial Early Classification
c. Marine →basic principles
i. Intertidal – between physical
1. Aristotelian System
limits of high and low tide
-plants and animals
and thus exposed to air
-animals with and without blood
periodically
-animals on land and water
ii. Subtidal – below low tide
→similarities
line; exposed to air only
under extreme conditions; 2. Linnaean System
offshore area -still used nowadays
iii. Open-ocean – creatures *hierarchical – graded series; ranked; inclusivity
found in the pelagic zone; →organization
deep ocean *binomial nomenclature – genus + species 3.
2. Locomotion Strickland Code
a. Mobile – capable of locomotion -guideline for naming
b. Sessile – attached to substrate -British Association for the Advancement of Science
c. Sedentary – limited locomotory abilities
d. Planktonic – forced to drift or wander 3. Mode Modern Classification
of Nutrition 1. ICZN – International Code for Zoological
a. Herbivores Nomenclature
b. Carnivores -based from Strickland Code
c. Suspension or filter feeders 2. Phylocode – International Code for Phylogenetic
d. Deposit feeders – substrate/sediment 4. Nomenclature
Symbioses -probe on phylogeny; rankless taxa but still
hierarchical

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Systematic Approaches to Phylogeny 1.
Phenetics/Taximetrics/Numerical Taxonomy -
measures characters and apply computer
algorithms to determine which groups are alike
-no phylogenetic assumptions
-classify organisms based on overall
similarity
-to prevent subjectivity
2. Polyphasic Approach
-consensus approach to bacterial systematics
-integrating all available data maximally -
goal: collect as much information as possible
in order to generate a useful classification
scheme
• Challenges
-enormous amounts of data
-large number of species
-data fusion/aggregation which will
demand efficient and centralized
data storage
→require collaborative efforts
3. Cladistics
-phylogenetic systematics
-science that concerns itself with discovery
and prediction of phylogeny
-emphasis is not upon the presence of all
shared traits, but upon the presence of shared
derived traits (synapomorphies)
→cladogram (visual representation of a
clade)
→parsimony (least number of evolutionary
transformations)

Criteria Used to Recognize Character

States 1. Ontogenetic criterion
2. Ingroup commonality
-similar = primitive
3. Stratigraphy
-fossils
-character in fossil = primitive
4. Biogeography
-more distant from place of ancestor, more
advanced
5. Function/Adaptive Value
-novel characters = specialized
6. Underlying synapomorphy

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The Poriferans and Placozoans enters this cavity
-not related but constitute simplest forms of metazoans 5. Ostium – narrow openings where water flows into the
spongocoel
Phylum Porifera 6. Osculum – larger openings where water exits the
-commonly known as sponges spongocoel
• Phylum Choanomonada/Choanozoa 7. Choanocyte
-“protozoa” or first animal a. Flagellum and Microvilli (Filter) – beating
creates a current for water to enter
General Characteristics b. Amoebocyte/Archeocyte – transport nutrients
-sessile and attached: feed on suspended particles -have to other cells of body and produce materials
nothing: no body systems for skeletal fibers (spicules)
-distribution: no terrestrial, 2% freshwater, 98% marine -functions:
-amorphous: no distinct body morphology but -generate currents that help maintain circulation
asymmetrical predominates of seawater within and through the sponge
-exhibit alloincompatibility: can distinguish its own cells -capture small food particles
and which cells are not of the same species -capture incoming sperm for fertilization
-with histocompability memory
-may lead to tissue disharmony
-indeterminate growth: enlargement without fixed upper
size limit
-filter feeders
-cell pigments and endosymbiosis (different colors)
• Importance
o Major component of aquatic ecosystem
o Provide shelter
• Histoincompatibility in Sponges
o Isogeneic Treatments
-fuse
-same colony
o Allogeneic Treatments •
-never fused Bauplan
-different colonies -microvillar collars surround flagella with units
arising from either single cells or syncytia
Parts • Specialized Cell Types
1. Mesohyl – gelatinous matrix that separates 2 layers of o Lophocytes – secrete collagen fibers from
cells in wall trailing end as they move at mesohyl
2. Epidermis/Pinacoderm/Pinacocyte – made up of o Spongocytes – secrete collagen that
contractile cells; outer layer polymerizes into “spongin;” only in Class
3. Porocyte – channels; entrance of water; doughnut- Demospongiae
shaped o Sclerocytes – secrete mineralized skeletal
4. Spongocoel/Atrium – water passing through porocytes spicules

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o Myocytes – musclelike cells aggregating

around oscula of some demosponges
• Spicule Formation
 structures present before regression
• Asexual Reproduction
o Gemmule formation
o Fragmentation
o Budding
• Sexual Reproduction
-hermaproditic: male – choanocytes, female –
archaeocytes
-internal fertilization
-most sponges “brood” the developing embryo
before released as swimming larvae while few
shed newly fertilized eggs

-importances: 2 Types of Larvae

-species identification 1. Amphiblastula
-support/maintain shape -hollow, swimming sponge larva
-discourage predation -flagellated at one end only
2. Stereoblastula
Gemmules -coeloblastula becomes solid as numerous cells
-dormant structures, typically far more resistant to detach from the wall of the blastula and
desiccation, freezing, anoxia completely fill the blastocoel
1. Archaeocytes phagocytose other cells to
accumulate nutrients
2. Cluster within the sponge
3. Cells surrounding each cluster will secrete a thick,
protective covering. The gemmule consists of the
cluster cells plus the surround medium ~ period of
vernalization, a period of dormancy,
developmental arrest ~ period of asexual
reproduction Type
4. In appropriate environmental conditions, gemmules
will hatch ~ can survive up to about 25 years
-results in identically the same individuals s of Body Architecture/Canal Systems -asconoid, syconoid,
leuconoid
• Adaptation → Pronounced Tissue Regression o
During unfavorable weather conditions
o Body becoming reduced to a compact
cellular mass with an outer protective
covering
o Cells reactivate when environmental
conditions improve, regenerating all of the
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 Class Calcarea
-spicules composed of CaCO3
Class Demospongiae
-spicules and fibers may be composed of spongin and/or
silica but never CaCO3
-may also contain chitin
-all freshwater sponges

Class Hexactinellida
-six-rayed spicules of silica and chitin
-aka glass sponges
-entire sponge is syncytial (having many nuclei contained
within a single plasma membrane) rather than cellular and
lacks contractile elements, thus no pinacoderm

Class Homoscleromorpha
-most homoscleromorh species lack spicules -when spicules
are present, they are entirely siliceous
-have a clear, distinct basal membrane underlying the
epithelium
-all epithelial cells bear cilia
Sponge Diversity
Read:
1. Phylum Placozoa
2. Introduction to the Hydrostatic Skeleton
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Phylum Cnidaria some extent locomotion

o Flowers of the Sea 3. Defense
-marine and some in freshwater 4. Species identification
-some colonial, only a few mm
-some may reach up to about 2m in diameter - o 3 Major Categories
Cyanea capillata/Lion’s Mane Jelly o Spirocyst – Volvents
-~10 000 species -for entangling prey
-sessile, they can only move feebly, fairly slow -thin-walled; undischarged tubule is
coiled like a spring
General Characteristics -discharged spirocyst lack barbs
-symmetry: radial -have minute sticky threads radiating
-mesoglea: gelatinous layer between epidermis and from wall of tubule like bristles
gastrodermis o Nematocyst – Penetrants
-contain living cells -most common
-epidermis -found in all groups of cnidarians
-gastrovascular cavity (GVC) -have spines or barbs on the surface of
-tentacles the everted tube (for puncturing)
-mouth/anus -inject toxins that paralyzes small
crustaceans
Bauplan -30 different types of nematocyst
-venomous cnidarian stings feel like a surprise o Glutinant
injection of needles -nematocyst characterized by an open,
-possess tentacles that bristle with tiny cnidae sticky tube used for anchoring the
cnidarian
Cnidae -secrete sticky substance that helps in
-produced by cnidocyte (cnidoblast/nematoblast) - fastening the tentacles to solid objects
have cnidocytes in localized parts of gastrodermis to and help in food capture and locomotion
quell swallowed prey -nematocyst: most common
cnidae (~synonomous) Life Cycle of a Jellyfish
-3 milliseconds

Cnida Structure
1. Has an opening/aperture
2. Often occluded by a hinged operculum 3. Consists
of a rounded, proteinaceous capsule: thickened and
stiffened by collagen
4. With sensory cilium: mechanoreceptive and
chemoreceptive
~Cnidocil: stiff Body Plan
5. Within a sac is a long, hollow coiled tube: everted 1. Medusa – resembles a gelatinous saucer or
upside down cup, jellyfish form, free
Function of Cnidae swimming/floating, mouth is usually in concave
side
1. Wrapping around small objects – food, collection,
prey 2. Polyp – tubular body, hydroid form, stationary;
2. Penetrating surfaces – food collection, prey; at resembles flower and stalk

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Nervous System elongated cells with contractile bases
-cnidarians possess bonafide nerves and muscles -a -either form longitudinal or circular musculature
central nervous system is lacking -movement of contractile cells may facilitate
-neurons and neurites passage of food
-surface area affected by stimulation of a cnidarian 1. Longitudinal: running from base to the tip of the
nerve cell varies directly with the frequency of tentacle
distribution 2. Circular: running around circumference of a body
column
• Interstitial Cells/I-cells
-multipotent cells that originate from embryonic
endoderm but migrate to 3 layers of body
-can be become neurons, gland cells, gametes, and
cnidocytes, and sclerocytes (anthozoans that
secrete spicules)

Diet and Digestion

-primary carnivorous
-soft corals: ingest phytoplanktons
-obtain nutrients from endosymbionts (algae):
characteristic of reef-building (hermatypic) corals Respiratory System
-lack gills and other specialized respiratory structures
Musculature -gases diffuse across all exposed epidermal and
-cnidarians are diploblastic: no true muscularis layer gastrodermal surfaces
-but they have muscle cells that are:
-found in their epidermis and gastrodermis Subphylum Medusozoa
-anchored to the mesoglea -recently established
1. Epitheliomuscular cells: ectodermal 2. -all cnidarians except Anthozoa
Nutritive-muscular cells: endodermal - -mitochondrial DNA genome is linear rather than
circular

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Class Scyphozoa -cup animals, with consistency of jelly hence the

-real jellyfishes; collectively known as jellyfish/sea name
jellies -few hundred species, all of which are marine and
many of which are quite large (up to 2cm across) - vascular = circulate O2 and CO2
jellyfish morphology is medusoid. The body is in -obtain nutrients as predators and from
the form of an inverted cup, with nematocyst studded zooxanthellae (unicellular algae)
tentacles extending downward from the cup, or bell.
The mouth is borne of the end of a
muscular cylinder known as the manubrium -bell
contracts → water expelled downward → animal
propelled to opposite direction
-mesoglea acts as a skeleton, stretching the
contracted muscles when they relax

Sophisticated Structures
-with sensory receptors
• Rhopalia
-club-shaped structures
-associated with dense aggregation of nerve
tissues
-act as pacemakers, triggering the rhythmic
contraction of the swimming bell
-distributed along the margins of the swimming
Digestion bell
1. Food particles captured by the nematocyst at the • Ocelli
tentacles -simple light receptors
2. Ingested at the mouth and conveyed to the -with pigments
stomach via the manubrium -non-image formers
3. Food is then distributed in 4 gastric pouches: with • Statocysts
gastric filaments: nematocyst-bearing tentacles -balance organs
that secrete an array of digestive enzymes -rhopalian-statocysts system provide a
4. Food is moved in the gastrovascular cavity mechanism through which the animal can be
(GVC) through the presence of cilia, and through informed of its physical orientation, whether
muscular contractions tilted or horizontal
-for organisms of this class, gastric cavity is the • Sensory lappets
GVC -touch/chemical receptors
-gastro = function in digesting materials -

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 genet; independent but genetically identical unit of
a genet

Sophisticated Behaviors
-periodic vertical migrations from surface waters to
deep waters and back again
-formation of temporary breeding aggregations -
strobilation
-transverse division of body into segments
which develop into separate individuals
-forms numerous individuals stacked on top
of each other
• Gonochoristic
-gono – reproductive organs
-chorist – separate Class Cubozoa
• Dioecious Defining Characteristics
-2 houses -medusa with boxlike body
-with few exceptions, individual medusae are -rhopalia bear complex, lensed eyes
either male of female -all members of the class have a cuboidal
swimming bell
Life Cycle of Aurelia aurita -members (~25 species) are called cubomedusae -
1. Planula larvae are the results of union between medusa stage dominates the life cycle -typically
sperm and egg only a few cm in its largest dimension and
-in the form of highly ciliated, microscopic extremely transparent
sausage -4 tentacles or 4 clusters of tentacles
-non-feeding stage that attaches to substrate and -only 2-3 cm in diameter with tentacles over 30 cm
transforms into small polyp (scyphistoma) long
2. Scyphistoma is sessile and lacks ocelli and -tentacles boast highly virulent nematocysts
statocysts →earned the title “sea wasps”
-feeding stage -eat small fish, annelids, and arthropod crustaceans,
-reproduces asexually by budding often killing individuals much larger than
-during strobilation, body divides transversely, themselves
forming numerous modules stacked on top of -some species can kill humans, inflicting pain -
each other intense pain
-each module eventually breaks from the stack as a -cardiac dysfunction
swimming ephyra (grows and gradually becomes an -respiratory depression
adult scypozoa) -rash lasts for several months
-genet: single fertilized oocyte producing a single -possess an unusually well-developed nervous
genotype system and remarkably complex eyes that can
-ramet: individual organisms that arise from a probably form images

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-the swimming bell curves inward at the lower edge, expelled when the bell contracts. This increases the force
restricting the size of the opening through which water is with which the water exits the bell, and thus the amount
of jet propulsion obtained. The effect is: -usually possess a shelf of tissue (the velum)
-greater propulsive force results from pushing the extending from the edge of the swimming bell toward
same volume of water out through a smaller the manubrium
opening in the same amount of time -the velum causes water to be ejected from under the
-cubozoans also differ from the true jellying in that the swimming bell through a narrower opening, and thus
polyp stage does NOT STROBILATE. The polyp with greater velocity, the effect is identical to that of
resulting from a single planula larva buds off more cubomedusae
polyps, each of which develops into a single medusa *scyphozoan medusae lack a velum

• Larva
-the planula larvae of hydrozoans develop from
Class Hydrozoa fertilized eggs, and the planula typically
-characterized by greater representation of the polyp metamorphoses into a sessile polypoid individual
morph in the life cycle lacking both statocysts and ocelli
-medusa morph dominates in a few members - -in Hydra, each polyp is separate, distinct being,
gastrodermal tissue lacks nematocysts: restricted to completely responsible for its own welfare
epidermis -some species harbor unicellular green algae, the
-no cells found within mesoglea zoochlorellae, in tissues
-more than 3000 species -lacks a medusa stage
-mostly marine -most other hydrozoans are colonial in the polyp
stage – a single planula gives rise to a large number
Subclass Hydroidolina of polyps, called zooids (or modules of a colony), all
• Medusa of which are interconnected and share a continuous
-mesoglea layer of the hydrozoan medusan is thick GVC
-mouth is borne at the end of a manubrium -ocelli -the zooids are often connected to each other, or to a
and statocysts are present substrate, by means of a rootlike stolon. The oral end
-all medusae are gonochoristic, a given individual of polyp (i.e. the end bearing the mouth and tentacles)
being either male or female but never both is called the hydranth

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-the stolons and stalks are commonly encased in a

transparent protective tube, known as perisarc,
composed of polysaccharide, protein, and chitin
-perisarc surrounding the hydranth is known as a
hydrotheca, and the hydroid is said to be thecate
(as opposed to athecate)
 -medusae of some species can “reverse” their
ontogeny under stressful conditions
o Dimorphic – 2 types of modules
o Polymorphic – more than 2 types of modules •
Dactylozooids
-for defense and feeding and reproduction,
heavily studded with nematocysts
-many form species-specific symbiotic associations
with other animals, including fish, sea urchins, sea
squirts (ascidians), polychaete annelids, gastropods,
bivalves, crustaceans, bryozoans, sponges, sea
anemones, and even other hydrozoans

Order Siphonophora
→carnivores
-free-floating hydrozoan colonies in which medusoid
and polypoid morphs are present simultaneously in a
number of different incarnations
• • Modified Medusa
Gastrozooids
o Pneumatophore
-modules specialized for feeding
-gas-filled float
-collect small animals using tentacles (with
-mesoglea reduced or absent
nematocysts) and ingest the prey through GVC
o Nectophore
-digestion is extracellular in the GVC, and then
becomes intracellular as the partially digested -modified to propel the colony
food is distributed throughout the colony through water by jet propulsion
• Gonozooids -lack mouth and tentacles
-specialized for producing medusoids and must • Modified Polyp
depend upon other members of colony for o Bract
nutrition; lack tentacles and are incapable of -leaflike defensive modules
feeding -aka phyllozooids

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-numerous nematocysts
 Myxozoans
-spore-forming parasites classified until recently as
protozoans
-infect annelids, bryozoans, and fish
-thought to be a peculiar, degenerate bryozoan, but it
turns out that the DNA samples leading to that idea had
been contaminated
-infective sporozoites emerge from the spore following
rupture along the suture
•
*Polar Filament
-must actually be a nematocyst (ultrastructural)
Muggiaea
-lacks pneumatophore
-gonozooids, dactylozooids, phyllozooids,
gastrozooids
-nematocyst-bearing structures on tentacles of
gastrozooids
• Physalia
-lacks nectophores, moved by wind and water

Class Staurozoa
-about 4cm long
-cold, shallow waters, attached to seaweeds, rocks, and
gravel by a stalk
-feed mostly on small crustaceans, catching prey with
eight sets of tentacles at their distal end -tentacles end in
great clusters of nematocysts • Life Cycle
-live the life of polyps
-no swimming medusa stage
-release gametes, planulae are not ciliated (creep over
substrate by muscle contractions) -no swimming stage
in whole life cycle. Crawling larvae eventually attach
to a substrate and soon develop feeding tentacles and
gonads
Order Hydrocorallina
-hydrocorals
-colonial
-largely restricted to warm waters
-not true corals
-dactylozooids abundant
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Subphylum Anthozoa Defining Characteristics

-flower animals -absence of medusa stage
-sea anemone, sea pansy, sea fan, sea whip, corals - -absence of operculum and cnidocil
exclusively marine -mitochondrial DNA is circular
-either solitary or colonial -presence of siphonoglyph
-coelenteron partitioned by mesenteries

Sexual Reproduction
-lacking a medusa, gametes are produced directly by
the polyp. A planula larva develops from the fertilized
egg and metamorphoses to form another polyp
-gonochoristic
-sequential hermaphrodism

Asexual Reproduction
-longitudinal/transverse fission
-pedal laceration: parts of pedal disc (foot) detach from
the rest of the animal and gradually differentiate to
form a new single-module ramet -fragmentation
resulting from violent storms can also lead to
substantial asexual replication

Anthozoan Morphology
-primarily carnivores
• Siphonoglyph
-ciliated groove in the pharyngeal wall leading
from the mouth
• Mesenteries/Septa
-distinct sheet of tissues partitioning the
coelenteron
-greatly increase the surface area available for
secreting digestive enzymes and absorbing
nutrients
• Acontia
-loaded with nematocysts and secretory cells and
can be extended outside the body through small
pores in the body wall. Used both offensively and
defensively, and they may also function in
digestion
• Acrorhagi
-covered with plant nematocysts; used in
defending a territory against invasion by other
anemones, even different genets of same species
-lack acrorhagi: catch tentacles, for aggressive encounters
among individuals

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Subclass Hexacorallia CO2, and nitrogenous wastes essential for algal

-possess many tentacles around the mouth opening and 6 growth
pairs of primary mesenteries -solitary: sea anemones -protected from herbivores
-colonial -occur in tissues at concentrations of ~1 to 5
-scleractinian corals (Order Scleractinia) -true (or million cells per cm2
stony) corals, which secrete substantial external Subclass Octacorallia
CaCO3 skeletons -8 complete mesenteries and 8 tentacles -tentacles are
-reef-building (hermatypic) or pinnate: bear numerous outfoldings known as pinnules
not (ahermatypic) -some ingest phytoplankton
-warm waters/tropics -all form modular colonies
-Great Barrier reef Australia: -anti-cancer agents
2000 km deep -single siphonoglyph
-get food through symbionts (algae)
-zooxanthellae have access to wastes of coral,
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PHYLUM CTENOPHORA Greek:

• Superficially resemble some jellyfishes and have
anatomical similarities similar with them
comb-bearing • Radially symmetrical: the main axis of the body is
oral-aboral
• Superficially resemble some jellyfishes • Most are
near-transparent or they lack in color, 3 Body layers
• planktonic 1. Outer epidermis: bilayered
• some benthic organisms are pigmented and use -Outer layer consist of multiciliated cells, mucus-
color patterns to camouflage with their secreting cells (secrete protective mucus covering
environment the body) –collocytes
• 150 species described 2. Inner layer
-consist of myoepithelial cells and nerve net
DEFINING CHARACTERISTICS - gastrodermis
1. Plates of cilia arranged in rows ~ ctene (~comb) 3. Middle mesoglea
2. Adhesive prey-capturing cell -thick and buoyant; houses nerve, pigment and
amoeboid cells (secrete and maintain the matrix of
GENERAL CHARACTERISTICS the mesoglea)
CN: Sea walnuts, comb jellies: ctene~comb • All Bioluminescent: exhibited by nearly all ctenophores;
are marine results from a chemical reaction in which much of the
• Nearly all are predators (of larval fishes and their excess energy is given off as light rather than as heat
eggs); only 1 parasitic is known • Almost all are Iridescent: colors are generated by the diffraction of
described as planktonic (float about in their medium), incident light (of the comb rows)
carried about by ocean currents Functions: mate location, species recognition, luring of
• All are predators of zooplankton: such as prey, startling of predators
crustaceans, jellyfish, fish eggs, and often each
other ~ sea pests DIGESTION AND INTERNAL TRANSPORT Carnivorous
• Play a major role in estuarine/marine ecology – by organisms feeding mainly on crustaceans and other jellies
influencing the success of commercial fishes 1. Tentacles: originate on the aboral hemisphere;
• Diploblastic like cnidarians Tentilla: lateral row of thread-like filaments, filled
 with adhesive granules. • Aboral organ located in the aboral end of the
2. Mouth: direct seizure with the mouth; or may use ctenophore
tentilla or tentacles. • rudimentary (one of the earliest) brain 1.
3. Stomodeum: (pharynx) begins the extracellular Statocyst
digestion. 2. Statolith
4. Stomach: food is distributed in ciliary flow to all the 3. Balancers
digestive canals. FUNCTIONS:
5. Digestive canals: where particles flow until they are • Controls the locomotory comb rows – coordinating
endocytosed by nutritive cells, in which they the activity of the ctenes • Photoreception
undergo intracellular digestion • Chemoreception
• 2 end as blind sacs • Pressure/mechanoreception
• 2 are continuous with the anal
pores REPRODUCTION AND DEVELOPMENT • Clonal
6. Anal pores: where rejected particles leave the body (asexual) reproduction is not common, occurs only in
benthic platyctenids, which can excise fragments from
NERVOUS SYSTEM the margin of their flattened bodies.
• Apical sense organ

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• All are hermaphrodites except the 2 species of CLASSIFICATION

Ocyropsis (gonochoristic: produce only either • Based on the presence or absence of tentacles
male or female gametes)
• In response to light cues, gametes are spawned and Class Tentaculata
the eggs are fertilized in seawater through Class Nuda
External fertilization (except in some platyctene • For a small taxon, these organisms are remarkably
that fertilize eggs internally) diverse
• Cleavage is determinate: cell fates are fixed at the
first cell division A. Class Tentaculata
• Gastrulation occurs by: • Long, retractile tentacles
Invagination: groups of cells push into the • “spin-capture” feeding
blastocoelic space • Planktonic spherical/oval ctenophores • 2 well-
Epiboly: sheets of micromeres spread overs over developed tentacles with tentilla
what were the adjacent macromeres
• Embryonic development is direct and leads to i. Order Cydippida
cydippid larva: planktonic juvenile • Long, retractile tentacles
• Planktonic spherical or oval ctenophores
Gastrulation • Usually 2 tentacles that are well developed and
Delamination: cells of the blastula divide into the with tentilla (side branches of the tentacles)
blastocoel, forming an inner and outer cell layer, between ii. Order Lobata
which the mesoglea is later secreted Ingression: certain • Common planktonic comb jellies;
cells become detached from their neighbors, and simply • Body somewhat compressed laterally – only 4 of the
move into the blastocoel, creating a 2nd layer of cells comb rows are fully developed
• Reduced tentacles but have 2 muscular oral lobes
• Oral lobes covered with mucus and colloblasts and
MUSCULATURE act as tentacles that constitute the primary food
• Muscles are derived from AMOEBOID CELLS within collection surface; may also use in locomotion
the mesoglea thus resides in mesoglea layer when muscular contractions happen
• Have genuine smooth muscle tissue • 1st known • Only 4 comb rows
smooth muscle fibers have been isolated from • Auricles: bears tentilla (side branches with lots of
Mnemiopsis leidyi and Beroe sp. colloblast); assist in prey capture
• Plays little or no direct role in locomotion • May be
used in escaping predators iii. Order Cestida
• Body compressed laterally forming long ribbon; with
 the mouth and the apical sense organ on opposite • Body forms flattened plate and have a different plane
sides at its midpoint such that the oral and aboral surfaces have moved
• Locomotion thru 4 comb rows; sinuous muscular toward each other
movement and sometimes movement of cilia • Comb rows reduced or absent (PRESENT IN
• Feeding accomplished not through the ribbon but EARLIER STAGES OF LIFE)
through the short tentacles in the oral region • Locomotion: Floaters or gliders; creeping slowly over
solid substrates using muscular contractions of the
iv. Order Platyctenida body or ciliary movement
• Adults bear 2 long tentacles

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v. Order Beroida
• Lack tentacles and oral lobes: such that
feeding is through the muscular mouth
• All 8 comb rows/costae well developed
• With macrocilia inside mouth, consisting
of thousands of axonemes, are located
inside the mouth; and are used as teeth
that can chop of large prey into bite-sized
pieces

General Morpho-anatomy of Ctenophore (a) and

ctene (b)
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Phylum Platyhelminthes through a single opening.

• Gas exchange is accomplished by simple diffusion
General Characteristics across the body surface; by being flat they have a
• 34,000 described species, with—at present—no relatively high surface area to volume to ratio such that
uniquely defining characters(synapomorphies), 80% sufficient gas exchange can occur to support an active
parasites • includes one class of mostly free-living lifestyle
individuals (the turbellarians) and three classes of
exclusively parasitic individuals (the monogeneans, Excretion
trematodes, and cestodes) • Leading view of evolutionary • Eliminate nitrogen from protein metabolism in the
relationships among the major platyhelminth group form of ammonia, which diffuses across the body
• Systematics is still much debated and decidedly surface
unsettled • Protonephridia excrete excess water and metabolites
including ions
• Metabolites waste move out of the body by diffusing
across the body surface OR… • They may have a series of
specialized organs known as PROTONEPHRIDIA: first
kidney
• Take the form of the MESH CUP → with cilia or
flagella (fluid is ultrafiltered) → convoluted tubule →
excretory pore

• All flatworms are acoelomate, triploblastic, and

bilaterally symmetrical. • most have no anus; food enters,
and unmetabolized wastes leave the digestive system,
 Functions:
1. Play a role in osmoregulation 2. Eliminate
metabolic wastes
Flame Cell: bunch of cilia that expel wastes to the
outside; the actions of the cilia somewhat resemble a
flickering flame
Solenocyte: single flagellum found within the cup

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Reproduction
• The vast majority of flatworm species, in all four
classes, are simultaneous hermaphrodites; each individual
can, at any one time, function as both a female and a
male. In consequence, sperm exchange and egg
fertilization can occur when any individual encounters
another individual of the same species. Individuals • bear
generally cannot fertilize themselves.
one or more pairs of eyes anteriorly, along with a variety of
Class Turbellaria cells that sense chemicals (such as potential foods),
General Characteristics pressure changes (such
• Most of the 4,500 turbellarian species are free-living, as those produced by water currents), and mechanical
but about 150 species are commensal or parasitic with stimuli.
other invertebrates. • individuals in about 10% of the species also have
• Mostly marine; few freshwater; few species are statocysts.
considered terrestrial, restricted to very humid areas. Habitat
• Typically less than 1 cm long • Most aquatic turbellarian species are benthic; that is,
they live in or on the ocean, lake,pond, or river bottom.
Nervous System
• possession of a cerebral ganglion—a primitive but Locomotion
distinct brain— and from one to three (rarely four) pairs • The body’s outer surface is ciliated, often more so on
of longitudinal nerve cords (Fig. 8.3). Such an advanced the ventral surface than on the dorsal surface.
nervous system also characterizes the parasitic members • Most species move at least partly by secreting mucus
of the phylum from the ventral surface and beating the ventral cilia
within this viscous mucus. • an increased number of cilia
in contact with the substrate compensates for the
increased weight of a larger animal, and the ability to
move need not suffer as the animal grows.
• In contrast to the monociliated condition of cnidarians
and sponges, each flatworm epidermal cell is
multiciliated, bearing several to many cilia.
 attaches posterior, releases anterior, organism moves
forward

1. Pedal waves: subtle waves of muscular contraction

along the animal’s ventral surface; unidirectional, moves
from anterior to posterior

2. Looping: anterior attaches, pulls posterior forward,

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The Platyhelminths 151
seems to produce a viscous glue,
For effective looping locomotion, the flatworm must be
able to adhere locally to the substrate to prevent sliding 4. Frontal gland: an anterior aggregation of secretory cells
backward while the pulling and that is a characteristic of most turbellarians; roles in
(duo-glands)
defense, slime production for locomotion and adhesion
pushing forces are being generated. On the other hand,
attachments to the substrate must be only temporary if the
animal is to progress forward. Flatworms typically Digestive System and Nutrition
possess a large number of , • Incomplete digestive system: they have a mouth but
paired secretory cells (duo-glands) located on the ventral don’t have an anus, it’s a blind sac. Mouth serves as the
surface and opening to the exterior. One cell of each pair anus also. – blind-ended gut
Figure 8.5 rhabdites
Diagrammatic illustration of looping by turbellarians. Once an rhabdoids

while the other cell presumably secretes a and composed of phagocytic and gland cells. Gut is
attachment to the substrate is made at point X (a), contraction of
the longitudinal musculature brings the rear of the animal anteri
orly (b,c). The anterior attachment is released, and a new attach-
ciliated.
chemical that breaks this attachment to the • Shape of the gut: determined by the size of the worm
ment is made at the posterior end (d) at point Y, allowing the head
of the animal to be thrust forward (e,f).

substrate (Fig. 8.6).

rhabditeone duo-gland

ventral
surface of
turbellarian

releaser gland

adhesive gland

sensory cilium

substrate

Figure 8.6

• Wall of the gut is a simple epithelium (single layered)

Diagrammatic illustration of the ventral surface of a free-living turbellarian flatworm, showing the arrangement of the duo-gland system. Adhesive glands that dissolves the attachment as appropriate. Rodlike rhabdites, as illustrated, are encountered in the epidermis of most turbellarians; a defensive
produce a chemical that attaches part of the animal to a substrate; the releaser glands secrete a chemical function has been suggested.
Modified from Tyler. 1976. Zoomorphology 84:1.

1. Duo-glands: paired secretory cells; glandular adhesive 2. Rhabdoid- rod shaped, membrane bounded secretion that
organs that is used for temporary adhesion in
pec24182_ch08_147-178.indd 151 17/12/13 12:59 PM expands to form mucus that coats the animal’s body in
interstitial organisms (that live in between the spaces of sand response to avoid predation or to avoid dessication
grains); composed of 2 glands –
• viscid gland that secretes adhesives and cements the animal 3. Rhabdite: small and cylindrical aggregations of cells that
to the substratum, and the secrete a thick mucous that coats the animal’s body, possibly
• releasing gland that secretes the de adhesive, the substance in response to attempted predation or dessication
that breaks the attachment
• Microturbellarian- simple and unbranched enzymes <proteolytic enzymes> and endopeptidases of gland
• Macroturbellarian- gut is branched • Order Acoela: lacks a cells) • Intracellular digestion is by phagocytic gut cells
gut cavity; after cells that are multinucleated will be cells that • Turbellarians can withstand prolonged starvation by
secrete digestive enzymes reabsorption of systemic tissues including the parenchyma
tissue.
• Carnivorous; Feed on a variety of invertebrates • Capture prey by wrapping themselves around it, entangling
• Digestion is first extracellular (thru actions of pharyngeal it in slime or by pinning it to a substratum by means of their
adhesive organs.

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Some paralyze prey with their toxic mucus, while others routinely by asexual fission, an ability closely linked
use their penis to stab their prey. • Food maybe swallowed to their great regenerative capabilities
whole or in pieces. • It is not yet clear why the members of some
• Some simple organisms especially the ones without the turbellarian species can regenerate while others
gut absorb nutrients through their body wall. cannot

• The mouth of more advanced species is often borne at

the end of a protrusible pharynx; other species
possess a separate proboscis that spears food items
and then transfers them to an adjacent mouth opening

Class Trematoda
-parasitic
-commonly known as flukes (leaning more on Digenea)
-leaf-like in structure
• For most other turbellarians, copulation may occur by -blind-ended gut (bi-lobed) → bifurcating -
hypodermic impregnation, in which the stylets of the
alimentary canal ends in a closed space -never
penis pierce the body of the
addition, many turbellarian species reproduce segmented

partner a protective capsule; in most turbellarian Neodermata

species, there is -new skin
• The eggs of each animal are released
-larval epidermis is replaced by non-ciliated
after fertilization and generally develop Turbellaria Trematoda Monogenea syncytial epidermis
directly into miniature flatworms within Cestoda -parasitic

no free-living larval stage in the life cycle • In several Subclass Digenea

marine species, however, the developing embryo gives -di: two
rise to a short-lived, microscopic free-swimming larval -genos: races
stage, -in its characteristic life cycle, it needs 2
IH – necessary for the life cycle of parasite; never reaches sexual maturity
-flukes DH – where sexual maturity [adult] of parasite is reached

-endoparasites
-2 or more hosts and at least 2 infective
stages -1st IH: gastropod mollusc
-2nd IH: arthropod/fish
-DH: vertebrate
-dorsoventrally flattened
• Regeneration is accomplished through
the activities of neoblasts,
undifferentiated cells – unique to
turbellarians – with remarkably versatile
developmental plasticity. In
-oral sucker → anterior-most -ventral
sucker/acetabulum
for attachment (formation of negative pressure)
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Life Cycle
1. Egg
-resistant
2. Miracidium
-ciliated, swimming larva
-apical gland (produce proteolytic enzyme)
-penetrate through body of intermediate host 3.
Sporocyst
-sack-like
-gutless
-no feeding apparatus
-specialized for production of new
individuals
*germ balls/germ cells – have the capacity to
develop into a new individual
4. Redia [Rediae]
-mouth, pharynx, gut, and embryo
5. Cercaria
-like a fish
-bifurcated end at the tail
-specialized to look for either next IH or
DH -have proteolytic enzymes
-have chemoreceptors
• Metacercaria
-encyst larvae; tissues of 2nd IH; excyst and
migrate to their characteristic location and
mature

Clonorchis sinensis
-Chinese liver fluke
-causes Chlonorchiasis
-epidemiology: high prevalence in East Asia
(Japan, Korea, China)
-life cycle:

-treatment: Chloroquine in Praziquantel

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-diagnosis: stool analysis -migrating juveniles in ectopic [abnormal] locations such
-prophylaxis: cook food thoroughly, sanitation, control of as the eyes, brain, skin, and lungs (rare)
IHs
Halzoun
Fasciola hepatica -buccopharyngeal infection
-caused by adult stage
-in Middle East, raw liver is eaten

Schistosoma japonicum
-sheep liver fluke -Asian blood fluke
-IH: Lymnaea sp. -causes Schistosomiasis
-causes Fascioliasis -spine determines sex
-Fascioliasis hepatica -S. haemotobium [Africa], japonicum [East Asia],
-Fascioliasis gigantica mansoni [Africa, South America, West Indies],
-sheep liver rot intercalatum [Central Africa]
-epidemiology: prevalent in agricultural areas, riverbeds, -epidemiology: prevalent in East Asia, damaged visceral
freshwater ecosystems (attack ruminants) organs
-life cycle: -IH: Biomphalaria sp.
-life cycle:

*zoonoti
-
c disease
-diagnosis: stool analysis
treatment: Praziquantel
-treatment: Bithionol, Emetine HCl, Praziquantel -
prophylaxis: clean properly, cook properly, control IH -diagnosis: muscle biopsy, stool analysis, blood sampling
-prophylaxis: precautionary measures in swimming
Liver Rot *ascites
-necrosis (dead tissue of liver) *swimmer’s itch
-anemia
-pipestem fibrosis of the bile ducts Dicrocoelium dentriticum
-cirrhosis of the liver -lancet liver fluke
-jaundice -causes Dicrocoeliasis
-gall bladder damage -epidemiology: prevalence in agricultural areas -life
cycle:

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Subclass Aspidogastrea
-80 species
-with large ventral sucker divided by septa
-bear resemblance with Monogenea and Digenea
-no haptor
 -also require an intermediate host
-reach adulthood in fishes and turtles

-diagnosis: stool analysis, immunodiagnostic test

-treatment: Praziquantel

Paragonimus westermanii Class Monogenea

-lung fluke -8000 parasitic species (25 000 are thought to exist)
-causes Paragonimiasis -most are ectoparasites
-greatly reduces on productivity and quality of life -1-20 mm in size
-etiology -haptor and prohaptor
-hosts: snails, crabs -haptor: attachment organ on posterior side
-mode of transmission: -prohaptor: attachment organ on anterior side
-ingestion of crabs (oral sucker)
-contamination with metacercaria
-signs and symptoms
-cough of long duration
-hemoptysis (coughing of blood)
-chest/back pain
-not responding to anti-TB medications -
diagnosis
-sputum [spit] examination; spinal tap - -no intermediate host
treatment
-life cycle
-Praziquantel
a. Adult attached on fish
-life cycle: b. Eggs
c. Oncomiracidium
d. Attachment to fish
-life cycle of Dactylogyrus vastator

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a. Attached on gills (high oxygen concentration, d. Attaches on fish, finds gills

blood supply)
-lays eggs Class Cestoda
b. Eggs dislodged on gills Characteristics
-hatches to oncomiracidium 1. Scolex
c. Oncomiracidium develops into adult -finds -anterior-most
next host 2. Proglottids
 -segmented/metameric -DH: mammals
3. Loss of digestive tract -epidemiology: prevalent in temperate regions
-compensated by: -life cycle:
-flat body
-microvilli → extensions
-increase surface area for more
absorption
-microtriches on tegument
-tapeworms
-non-ciliated, syncytial tegument
*hexaca

nth: oncosphere

-3 types of proglottids: Dipilydium caninum

-mature -host: domestic dogs and cats. Often occurs in children
-immature (wider than long) -pathology: rare
-gravid (longer than wide) -treatment: Praziquantel, Niclosamide -eggs:
egg packets
Reproduction -each segment with 2 sets of male and female
-sexual reproductive systems
-simultaneous hermaphrodite -genital pore on each side
-each proglottid has developed male and female -scolex has a retractable pointed rostellum with several
reproductive organs circles of rose-thorn-shaped hooks -life cycle:
-proglottid
-scatter eggs
-life cycle:
1. Egg
2. Oncosphere larva
3. Cysticercus
4. Adult
Hymenolepis sp.
• Hymenolepis nana
-dwarf tapeworm (20-40 mm)
-only human tapeworm that may not utilize an IH
-causes Hymenolepiasis
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Taenia solium system

-infection: -life cycle:
-Taeniasis (from adult)
-Cysticercosis (by cysticercus)
-general form:
-Cysticercosis cellulosae
(solium)
-Cysticercosis bovis (saginata)
-Neurocysticercosis
-presence of cysticercus within
the human central nervous
 -Cysticercosis: chest pain, muscle pain,
uncontrollable breathing impaired CNS
(movement, thought processes, slurred speech)
-treatment: Praziquantel
-prophylaxis: cook food thoroughly

Diphyllobothrium latum
-diphyllo: 2 leavs
-T. -bothrium: sucking slits (for attachment)
-fish tapeworm
-infection: Diphyllobothriasis
saginata vs T. solium -life cycle:
T. saginata T. solium

Scolex

No hooks on rostellum With hooks on rostellum

Cuboidal Circular

Proglottid

More uterine branches Less amount (7-13)

with lateral arms (15- -1st IH:
20)

copepod (freshwater crustacean) -epidemiology:

-etiology: ingestion of raw/undercooked meat - prevalence in Asian countries -pathogenesis: vague
abdominal discomfort, diarrhea
manifestations: vague abdominal discomfort, diarrhea
-treatment: Praziquantel
-diagnosis: stool analysis, muscle biopsy (Taeniasis)
-prophylaxis: cook food thoroughly
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The Nemerteans -no true heart, the blood vessels lack one-way valves. The
Defining Characteristics blood does not circulate unidirectionally. Instead, it ebbs
-muscular eversible proboscis housed in a fluid filled and flows erratically, propelled largely by muscular
cavity, the rhynchocoel contractions associated with routine movements of the
animal
Habitat -a few species have hemoglobin in their blood, but most
-shallow water environments, crawling, burrowing into species lack any such oxygen-carrying blood pigment
sediment, or lurking under stones, rocks, or mats of algae
-some in freshwater Excretory System
-same terrestrial-moist environments -same with flatworms – protonephridial excretory
-a variety live commensally with Arthropods and systems
Molluscs, but only a few are parasitic
Digestion
Locomotion -unlike flatworms, with an anterior mouth and a separate
-superficially resemble the free-living flatworms anus
(Turbellaria). Like the flatworms, are ciliated externally -digestive enzymes may be secreted in sequence as the
and secrete a mucus food is moved through the gut by ciliary activity
-a few species can swim by generating relatively violent -carnivores: annelids and crustaceans
waves of muscular contraction -unlike flatworms, possess a hollow, muscular proboscis
(inside rhynchocoel)
Body Plan -distinct from the digestive tract, floating in a separate,
-elongated, unsegmented, soft-bodied worms -like fluid-filled, tubular cavity called the rhynchocoel →
turbellarians, most are flattened dorsoventrally and alternate name: Rhynchocoela (G. snout cavity)
possess circular, longitufinal, and dorsoventral muscles
-typically form several to 20 cm, but in some cases as Prey Capture: Proboscis
long as 30 meters
-the proboscis can be shot out with explosive force →
-“Ribbon worm” → common name
different opening from mouth -when musculature
around the rhynchocoel contracts, the pressure within
Nervous System
-similar to turbellarian: the cerebral ganglia form a ring the rhynchocoel rises
anteriorly and give rise to a ladder-like arrangement of -in most species, the proboscis is everted through a
longitudinal nerves with lateral connectives proboscis pore that is distinct from the mouth -some
-equipped with chemoreceptors and species have proboscis and digestive tract have come to
share an opening
mechanoreceptors, located in specialized pits and
-the proboscis is armed with a piercing stylet -
grooves on the body surface, and sensory bristles -most
paralytic toxin
species also possess pigmented photoreceptors → “eyes” -prey may be harpooned directly
-most commonly, it is first wound around the prey, and is
Gas Exchange and Circulatory System - stabbed repeatedly with barb
diffusion -some coil and secrete a sticky mucus to help hold the
-true circulatory system – blood circulates throughout prey
body through well-defined, contractile vessels

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-if there is no stylet, captured food is usually
transferred to the mouth when proboscis is
retracted
-with styles: paralyze the prey, retracts proboscis,
then moves to the prey to eat
-in some species, prey are ingested whole,
particularly if they are vermiform (worm
shaped)

Classification
-2 major classes:
• Anopla
-lack stylets
-mouth is posterior to the brain
• Enopla
-with stylet (order Hoplonemertea)
-mouth anterior to the brain

Protection from Predators

-soft bodied, lack behavioral defenses, shells,
spines, and other protective hard parts -may
burrow
-have bacterial symbionts (Vibrio alginolyticus) -
synthesize TTX (tetrodotoxin) – powerful
neurotoxins that provide an excellent defense
against predators

Reproduction and Development

-gonochoristic (dioecious)
-few hermaphroditic species are protandric (as it
ages, each individual first becomes male and then
becomes female)
-pilidium larvae
-imaginal discs
-can also reproduce asexually and sexually

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Phylum Mollusca Collective term for odontophore radular assembly and
-L: soft complex musculature
• DC *old teeth are being worn down while new teeth are
o Dorsal epithelium forming mantle ▪ Secreted continually being formed and added onto the ribbon
calcareous spicules • Method of Feeding (ROME)
or 1 or more shells o Radula
o Radula o Odontophore
▪ Cuticular band of teeth in the o Mouth
esophagus o Esophagus
▪ Not present in bivalves • Different strategies for catching fish o Conus
▪ For feeding striatus
o Ventral body wall develops into a ▪ Through a lightning strike
locomotory/clinging foot cabal of toxins
• Body Plan ▪ Instantaneous
o 100 000 species (living) o Conus geographus
o 70 000 species (fossils) ▪ Uses net strategy
o Dissimilar looking organisms ▪ Nirvana cabal of toxins
▪ Most malleable in the animal ▪ Causing irreversible
kingdom neuromuscular paralysis
o No “typical” mollusc • Mantle Cavity
o Not all have shells consisting of CaCO3 o Houses the ctenida
• Shells of most molluscs (including gastropods and ▪ Gills
bivalves) ▪ Exit site for excretory,
o Periostracum – thin, outer organic layer digestive, reproductive
o Prismatic layer – thick, calcareous middle products
layer • Ctenidium
o Nacreous layer – thin, innermost calcareous o When present, may have purely respiratory
layer function or collection or sorting of food
o Mantle – secreted by specialized tissue • Osphradium
o Pearl – foreign particles that are trapped o Chemoreceptor/tactile receptor
between the mantle and inner surface o Located adjacent to ctenidium
• Radula
o Feeding structure Class Polyplacophora
o Consists of a firm ribbon, chitin, protein -Gk: many plate bearing
o Found in numerous rows of sharp chitinous -shell forms as a series of 7-8 separate plates -800
teeth species
• Radular sac -3-10 cm
o Produces ribbon -intertidal/on rocks
o Underlain by a supportive cartilage • -articulating plates are embedded from mantle -
aesthetes
Odotophore (tooth bearer)
-derived from mantle
o Supports the radular sac
-extending from holes of plates (light receptors)
• Buccal mass (cheek)/Odontophore complex o

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-pharyngeal/sugar glands o Shell – series of 8 overlapping and articulating

-release amylase-containing secretions into the plates covering the dorsal surface
stomach -multisectioned
• Body Plan →reason why the body can bend and
 conform to a wide variety of underlying Subclass Prosobranchia
substrate shapes -Gk: anterior gill
o Girdle – chiton’s thick lateral mantle -free-living and mobile
-mostly marine
Class Aplacophora -possesses well-developed shell, mantle cavity,
osphradium, radula
-Gk: not shell bearing
-foot bears a rigid disc of protein called operculum
• DC
o Cylindrical
Subclass Opisthobranchia/Nudibranchia -Gk:
o Vermiform body with the foot forming a
posterior gill
narrow keel -sea hares, sea slug, bubble shells
o 300 species in deep ocean -marine
o Few mm – cm -< 2000 species
o Unsegmented body with numerous calcareous -trends toward:
spines in outer cuticle -reduction of shell
o With radula -reduction of operculum
-limited torsion during embryogenesis -
Class Monoplacophora reduction of mantle cavity
-Gk: one shell bearing -reduction of ctenidia (replaced by cerata –
• DC feathery gills from dorsal surface)
o < 1 mm to 37 mm in size • Sea slugs
o 3-6 pairs of ctenidia o For defense
o Multiple pairs of pedal retractor muscles ▪ Use nematocysts usurped
o 6-7 pairs of nephridia from cnidaria
o 20 extant species ▪ Produce chemical defenses
o Marine at 2000 m depth housed by cerata
o Single, unhinged limpet-shaped shell o Movement
▪ Can swim in short spurts by
Class Gastropoda flapping lateral folds of foot
-Gk: stomach foot called parapodia
• DC ▪ Possess rhinophores in 2nd
o Visceral mass and nervous system become pair of tentacles located
twisted (90-180o) dorsally
▪ During embryonic stage • Sea hares
▪ Exhibited torsion o Gill plume
o Operculum o Tentacle with rhinophore
▪ Proteinaceous shield on the
foot Subclass Pulmonata
o Largest molluscan class -L: lung
o 60 000 species -intertidal zones and estuaries
o Marine, freshwater, terrestrial environment -terrestrial or freshwater

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-few with operculum cavity opens on right side

-long radula to keep herbivorous diet -mantle cavity is highly vascularized and function as
-head bears 2 tentacles lungs through an opening called pneumostome
-torsion limited to 90o -ctenidia is lacking, replaced by fold of mantle tissue near
-nervous system not greatly twisted and mantle pneumostome
Class Bivalvia (= Pelecypoda) deposited by the animal (Figure 12.22). The foot projects
Class Bi • valvia (= Pelecy • poda) (L: two valved [G: ventrally and anteriorly, in the direction of movement,
hatchet foot]) bī-val ́-vē-ah (pel-iss-ih-pō ́- dah) and the siphons, when present, project posteriorly.

Defining Characteristics • Distinct growth lines typically run parallel to the shell’s
1. Two-valved shell outer margins (Figure 12.23). The foot projects ventrally
2. Body flattened laterally and anteriorly, in the direction of movement, and the
The class Bivalvia contains more than 9,000 siphons, when present, project posteriorly.
contemporary species, including clams, scallops, mussels,
and oysters.

Major bivalve characteristics include

1. A hinged shell, the two sides (left and right “valves”—
valva = L: a folding door) of which are closed by one
or two adductor muscles; a springy ligament springs
the shell valves apart when the adductor muscles
relax;
2. Lateral compression of the body and foot; 3. Lack of The Molluscs 241

cephalization: virtual absence of a head and associated

sensory structures; 4. A spacious mantle cavity, relative
to that found in other molluscan classes;
5. A sedentary lifestyle; and
6. The absence of a radula/odontophore complex.
Figure 12.23
Bivalves are primarily marine, but about 10–15% of all Growth lines in the shell of the bivalve Arctica islandica. The age
of the shell can be determined by counting growth lines because

species occur in freshwater.

the patterns of shell growth are seasonal.The most recent growth
lines are near the outer margin of the shell valve.This individual
was estimated to be 149 years old.
© Douglas S. Jones, Florida Museum of Natural Histor y.

• No bivalves are terrestrial. Four basic gill types of Bivalves

1. Protobranch- Gills are attached to each other by
• Members of most species are suspension feeders, using interfilamental ciliary junctions, small and leaf
their gill cilia to drive water like. They have an unmodified appearance like
phytoplankton and other microscopic particles from the that of gastropods, chitons, and cephalopods,
seawater. and, one may assume, Hecionelloids. These occur
only the most primitive bivalve groups, the
• A conspicuous bulge in the shell is frequently seen on Palaeotaxodonta and Cryptodonta.
the dorsal surface, adjacent to the hinge. This bulge, the
umbo, is comprised of the earliest shell material 2. Filibranch – Gills linked by ciliary disc Figure 12.24

(a) A protobranch bivalve, Yoldia limatula, with its foot and the lower part of its shell buried in sediment. (b, c) Two collection.

through the mantle cavity and capture types of

Arrows indicate the path of water flow through the mantle cavity. The gills function primarily in gas exchange, not food

junctions. These form lamellar sheets of

protobranch gill, with the more primitive (ancestral) gill type on the left. The anterior end of the animal projects into the

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(a) After Meglitsch. (b, c) After Russell-Hunter.

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Zoology 111 – Invertebrate Zoology Jules Raposas

individual filaments in a "W" shape. They hang filibranch type, but with cross partitions laterally
downwards into the mantle cavity but have their joining the filaments to create water filled
terminal portions bent upwards. These cavities between them. This is the most advanced
characterize many of the Pteriomorpha. gill type, and also the most common, found in by
far the largest number of species of bivalves.
3. Eulamellibranch – Junctions made of tissue rather than
cilia. These have the same "W" shape as 4. Septibranch – Lack filaments and form muscular
 septum. These gills are only found in the
Poromyacea, a superfamily of rock borers (Order
Pholadomyoida), and are clearly an adaptation to
the unusual lifestyle of their owners. They run
transversely across the mantle cavity forming a
sort of partition that divides the mantle cavity (a) A
into upper and lower sections, and almost
enclosing the internal chamber, but maintains protobranch bivalve, Yoldia limatula, with its foot and the
only a small connection with the outer cavity. lower part of its shell buried in sediment.
• One pair of gills is present in the mantle cavity, with
one gill on either side of the body (Fig. 12.24b, c).

• Include

Yoldia and Nucula

Subclass Protobranchia (G: first gill) • Morphologically primitive state of bivalve
Defining Characteristics • Bipectinate gills
1) Gills small and resembling those of gastropods, • Deposit feeding
functioning primarily as gas exchange surfaces;
2) Food collected by long, thin, muscular extensions of Subclass Lamellibranchia (G: plate gill) • Include familiar
tissue surrounding the mouth (palp proboscides) bivalves such as clam, mussels, scallops
• entirely marine • Gills modified to collect suspended food particles
• all species live in soft substrates (Fig. 12.24a). • Secretion of attachment byssal threads by byssal or
byssus gland in the foot

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Zoology 111 – Invertebrate Zoology Jules Raposas

A lamellibranch clam dissected by removal of its left Piece of wood riddles with X-ray photo of a piece of wood-boring
valve. Note the relative sizes of the gill and the tiny labial bivalves containing shipworms
palp compared to those in the protobranch. The gills act
Subclass Septibranchia/ Anomalodesmata (G: fence
as respiratory organs as well as feeding organs.
gill/G: irregular ligament)
• Small group of carnivorous bivalves and on pieces of
“Shipworms” – wood boring lamellibranchs
decomposing animal tissue • All are in very deep marine
Teredora malleolus
water • Ctenidium highly modified, lacking filaments and
• With wood-storing caecum packed with symbiotic forming muscular septum perforated by ciliated openings
bacteria
• Feed as vacuum cleaners sucking on small crustaceans
• Pallets analogous to operculum • Shell used to
and annelids
excavate the burrow
• Stomach lined with chitin
• Small labial palps with no sorting function
• Very reduced style with no crystalline sac

Cuspidaria, a septibranch clam. Septibranchs are often

called "dipper shells"
 38 of 56
Zoology 111 – Invertebrate Zoology Jules Raposas Subclass Coloidea

Class Cephalopoda (G: Head foot) • Chromatophores (colored cells) overlay iridocytes
• ~600 extant species (extinct ammonites) • Fast- (reflective cells)
moving, active carnivores capable of complex • Photophores (light organs)
behavior • Most have ink sac discharged through anus
• Exclusively marine • Male octopi have hectocotylized arms
• Largest cephalopod is the giant squid Architeuthis
– weighing 1,000 kg and reach 18m with 5m
long tentacles
• Smallest cephalopod is < 2cm • Derivatives of the
molluscan foot are the siphon/funnel, arms (8) and
tentacles (2) • Possess two eyes with cornea, lens,
iris, diaphragm and retina thus focusable and image-
forming
• Closed circulatory system with systemic and
branchial hearts
• Locomotes by jet propulsion
• Chromatophores in contracted (left) and expanded
(right) forms
Expansions and contractions are mediated by
muscle elements in the skin and under direct
control from the brain

Architeuthis dux

Subclass Nautiloidea
Nautilus
• 5 or 6 species possessing true shell • Shell divided
by septa into series of compartments
• Septa penetrated by siphuncle (calcified tube with
enclosed strand of vascularized tissue)
• Cameral fluid in chambers
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Phylum Annelida -control the actions of each body segment • Giant

-L: annulus = ring Axons
-run to the longitudinal muscles
Defining Characteristic -used in rapid retreat
-presence of setae/chaetae
-setae: bristles Excretory System
-chaetae: chitin -metanephridia
-project outward from the epidermis to provide -open at both ends
traction -nephrostome:
-made of beta-chitin -kidney mouth; where water is
-flexible and tough drawn through action of cilia
-strengthened by sclerotized -convoluted tubule:
protein or inorganic material -salt, aa, water is reabsorbed
(CaCO3) -nephridiopore:
-form temporary attachment sites and prevent -where urine and other metabolic
backsliding during locomotion byproducts comes out
-bladder:
Morphology -temporary storage
-exhibit metamerism (metamere/somite) -septa: thin
sheets of tissue that separate each segment Digestive System
-prostomium: anterior most part of the body; -straight tube
eyes (ocelli) and sensory appendages (near -foregut
clitellum) -midgut
-pygidium: posterior most part -hindgut
-body wall -mesenteries
-for locomotion -supports the gut
-gas exchange -double layer of membrane on inner surface of
-cuticle body wall
-collagenous -mouth → pharynx → esophagus* → crop → gizzard →
-non-living intestine
-limited permeability to gas and
-*calciferous gland
water
-release calcium carbonate to rid the earthworm’s
-epidermis
body of excess calcium
-glandular
-food moves into the intestines as gland cells in the
-setae (anchored)
intestine release fluids to aid in the digestive process
-muscular layer
-intestinal wall contains blood vessels where the digested
-circular (outer)
food is absorbed and transported
-longitudinal (inner)
-oblique (to avoid aneurism)
Reproductive System
-asexual
Nervous System
-fragmentation, budding, fission
• Central Nervous System -sexual
-dorsal brain -gonochoristic
-ventral pair of longitudinal nerve cords • -segmental gonads and produced in peritoneal
Ganglia/Segmental Nerves tissue (not in gonads) and
-mass/accumulation of nerve cells
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Zoology 111 – Invertebrate Zoology Jules Raposas

released in coelomic cavities (where they mature) -longitudinal muscles antagonize longitudinal muscle on
-spawn gamete through metanephridia - the other side resulting in rapid-eel like movement
fertilization is external →Erythoe sp.
-trocophore larva
-not only in annelids (annelids, molluscs, etc.) Subclass Sedentaria
→ annelids + molluscs = -spend their entire lives in simple burrows in the
LOPHOTROCHOZOA sediment of in simple rigid tubes – they live sedentary
-small, translucent, free-swimming larva of lives
marine annelids and most groups of molluscs -tubes are made up of calcium carbonate, proteins, and
-spherical or pear-shaped and are girdled by a polysaccharides
ring of cilia -parapodia and acicula are greatly reduced
-produced by echiurans and sipunculans -not by
clitellata Reproduction and Development
-produced by many molluscs • Epitoky – partial/full transformation for reproduction
• Reproductive module that detach from posterior of
1. Prototroch atoke → comingle with other epitokes → discharge
-located around the equator of the animal their sperm and egg -segmentation proceeds
-anterior to the mouth posteriorly
-animal’s main locomotory organ
• Epitoke – result of epitoky; sexually mature being
2. Metatroch
• Atoke – anterior portion of the original animal
-in between 2 bands of cilia
3. Telotroch
-located posteriorly on what will become the Family Siboglinidae (~Phylum Pogonophora) • DC
terminal portion o Gut tissue forms an organ (trophosome) that
becomes filled
Diversity with chemosynthetic bacteria
Class Polychaeta (chemolithoautotrophic)
-70% of annelids o Segmentation at the opisthosoma
-predominantly marine • GC
-eyes and sensory appendages on prostomium -DC: o Trunk contains pair of uninterrupted coelom
parapodia with body wall surrounded
-paired lateral outfoldings of the body - by outer circular and inner
important in ID (high taxonomic value) -↑SA, longitudinal muscles
o Regions of cilia and papillae and girdles of
vascularized (function in gas exchange)
setae
-locomotory function
*Acicula: chitinous support rods
o Gonads and trophosome within coelomic
*Elytra: series of overlapping protective plates cavity
o Opisthosoma for digging and anchoring
Subclass Errantia
-active, mobile species Subfamily Perviata/Frenulata
-move through the action of the parapodia, which are -6-36 cm and < 1 mm wide
operated in complex patters as oars -tubes thin walled and open at both ends

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Zoology 111 – Invertebrate Zoology Jules Raposas

-tubes permeable to DOM (dissolved organic matter) Subfamily Obturata/Vestimentifera

-one coelomic compartment per segment of opisthosoma -2 m long and 25-40 mm wide
-first discovered in 1977 near Galapagos Island -thick • GC
tube walls acting as barrier for DOM uptake (more o 150 described in shallow waters
applicable with chemosynthetic bacteria) o Few mm to 50 cm with extreme sexual
-nutritional supplement by chemosynthetic bacteria at dimorphism
trophosome o Deposit feeders with highly muscular
-possess long collard vestimentum and obturaculum plug proboscis which can be extended 25x the
CO2 + 4H2S + O2 → [CH2O]n + 4S + 3H2O length of body
o No segmentation of metamerism in extant
Class Clitellata species (only in fossil records and larval
• DC segmented nervous system)
o Pronounced cylindrical glandular region o Papillae in the trunk and setae in the posterior
(clitellum) that plays important roles in end
reproduction • DC
▪ Segment 32-37 o Muscular organs (anal sacs) outpocketing
▪ Production of nutritive fluids from the rectum into the coelomic space
and production of cocoons bearing numerous funnels that discharge
o Permanent gonads coelomic fluid (and wastes?) through the
anus
Subclass Oligochaeta o Unique sex determination
-Gk: few setae • Unique sex determination of Bonelia viridis o When
-~3 500 species (10 000 species) B. viridis is in the egg stage, its sex is not yet determined.
-mostly freshwater and terrestrial; few marine -septa During the larval stage, if the individual comes in contact
dividing coelomic cavity into semi-isolated compartments with a female, then the larva will develop into a female.
-moves by peristaltic waves This is because the female proboscis produces a hormone
→“hearts” – contractive vessels (septal hearts) which stimulates the larva to develop into a male and
resides in the nephridia (up to 25 males in proboscis)
Subclass Hirudinea
-Gk: leech Sipuncula (Peanut worms)
-500-630 described species -Gk: little tube
-mostly freshwater and terrestrial; few marine -with • GC
anterior (fused 1-4 segments) and posterior suckers (fused o Body wall composition and general life
25-33 segments) history similar with annelids
-no septa; no setae → may be remnants - o About 350 species
moves by looping o Few mm to 1 m
→mutual sperm exchange (simultaneous o Lack setae and show no trace of segmentation
hermaphrodites) o All marine in shallow waters
Echiura o Mostly deposit-detritus feeders
-Bonelia viridis o Gonochoristic

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o Do not secrete tubes, just form

burrows or live in empty mollusc
shells or polychaete tubes; in rock
crevices, calcareous substrates of
coral reefs
• Body Parts
o Retractable introvert bearing
numerous mucus-covered tentacles
withdrawn into the body during
 ingestion
o Unsegmented trunk with papillae
• Sensory Organs
o Ocelli
o Nuchal organ – chemosensory organ
(ciliated pit, groove, fold, or slit)
• DC
o Anterior part of body forming
eversible and fully retractable
introvert , with mouth at its end
o Anterior tentacles connected to series
of muscular sacs (compensatory
sacs) that pump fluid into the
tentacles
o Multicellular bodies (urns) in the
coelomic fluid specialized for
accumulating wastes

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Zoology 111 – Invertebrate Zoology Jules Raposas

Arthropoda Bauplan
• Epidermis produces a segmented, jointed, • 85% of all animal species to date belong to
and hardened →sclerotized) chitinous phylum Arthropoda
exoskeleton, with intrinsic musculature - Insects, spiders, scorpions,
pseudoscorpions, centipedes, crabs,
between individual joints of appendages
lobsters, brine shrimps, copepods, and
• Complete loss of motile cilia in adult and barnacles are all arthropods
larval stages 1. Segmentation
 • Metameric - Other arthropods →except chelicerates &
- New segments arise during development proturans) have single uniramous pair
• Tagmatization of antennae such as Myriapoda
- Fusion and modification of different regions →millipedes & centipedes), Insecta
of the body for highly specialized →insects) and the extinct Tribolita
functions
→trilobolites)
- Two major arthropod groups →insecta,
- Functions vary →sensing touch, air
crustacea) have 3 distinct regions
motion, heat, vibration, olfaction,
i. Head
ii. Thorax gustation)
iii. Abdomen
- Lacks cilia in the larval stage 3. Segmented & jointed appendages - In primitive
condition, each segment has a pair of appendages
2. Cephalization a. Crustacean → for swimming
• Anterior tagma →head, cephalon or b. Praying mantis → capturing prey
cephalothorax) c. Grasshopper → jumping
- Compounds eyes →has many individual
units called ommatidia) 4. Body walls
- In contrast to the human eyes, compound - Exoskeletons
eyes break up the image before it reaches - Generally hard, external protective covering
the retina, so that each ommatidium - Molluscan shell primarily functions to
samples protect soft parts within
only a small part of the complete image - Arthropod integument aid in locomotory
• Antennae skeleton
- Generally paired appendages →one or - Exoskeleton is secreted by epidermal cells.
two pairs) in arthropods The epicuticle →outermost layer) is
- Crustaceans → present only in the first two generally waxy, composed of
segments of the head wherein the lipoprotein layer underlain by layers of
smallest pair is known as antennules and lipid.
the larger pair is biramous

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Water-impermeable, hence outer body for strength, non

surface cannot function for gas exchanges allergenic reaction & biodegradable
- Cuticle make arthropods resistant to properties
water loss by dehydration. Epicuticle is 1. Solubilized and reformed
thin →3% total thickness of into fibers for fabrics and
surgical sutures
exoskeleton)
2. Capsule for therapeutic
- Procuticle → largely of polysaccharide drugs
chitin in association with a number of 3. Substitute for plastic wrap
proteins 4. Non-digestible and
- binding abilities to
*chitin has been commercially produced organic and inorganic
 compounds →reduce iridescence
calories & cholesterol
uptake) 7. Setae
5. Candidate for removing - Chitinous projections of the exoskeleton
toxic organic and secreted by trichogen cell: maybe
inorganic compounds branched, plumose or in the form of
from drinking water and spines
sewage treatment
8. MOLTING/ ECDYSIS of exoskeleton and lining
5. Locomotion of the guts
• Sclerites - New cuticle is secreted before old one is
→division of cuticles are separated by shed
plates and connected via articular - Old cuticle is split by uptake of water or air
or increased blood pressure
membranes
- Both under neural and hormonal control
- Antagonistic contraction of flexor and
• Other neurohormonal controlled mechanisms
extensor muscles
in arthropods
- Resillin → substance or “animal rubber” in a. Regulation of reproductive cycle b.
wing joints and Regulation of body fluid osmotic regulation
jumping legs →stores energy upon and c. Migration of light-screening pigments in
compression and releases it efficiently) the eye
- Hemocoel → hydrostatic skeleton in d. Movement of pigment granules within
chromatophore cells leading to gradual
arachnids allowing them to extend their
changes in color
legs by increasing the blood pressure
9. Nervous system
6. Coloration - Single muscle fiber innervated as many as 5
• Brought by hemoglobin, chromatophores or by different types of neurons

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Zoology 111 – Invertebrate Zoology Jules Raposas

10. Circulatory system

- Open: with ostia, heart, arteries,
sinuses, blood; in some with
capillaries → closed)

11. Excretory system

- Malpighian tubules

12. Respiratory system

- Spiracles & Trachea
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Zoology 111 – Invertebrate Zoology Jules Raposas

Echinoderms →Gk: spine skin) Larva bilateral

and usually free swimming. Many readily
General characteristics regenerate body parts
• Sea lilies, feather stars, brittle stars •
Deuterostomes Water vascular system →WVS)
• 6 500 marine →living, few estuarine, 13 000 • Major unifying characteristic of Phylum
living fossils) Echinodermata
• Cephalization is lacking, body is designated as - Consists of fluid-filled canals derived from
oral & aboral coelomic compartments →hydrocoel)
• Lack circular muscles that form during embryonic
• Internal skeleton is 95% CaCO3 • 85 development
species are venomous - Hydraulic system
• Has ring canal → radial canal → ampullae →
Defining characteristics tube feet
1. Pentamerous radial symmetry in adults 2.
Calcareous ossicles forming an endoskeleton • Has ring canal → Stone canal → Medaporite
3. Water vascular system →WVS) servicing the • Has ring canal → Tiedemann’s body and
tube feet pollan vesicle →modified storage organ)
4. Ring canal & radial canal, exiting the tube feet
5. Mutable connective tissue that rapidly polymerize • Transmission system uses pressurized
from stiff and hard to soft gel & vice versa hydraulic →relating to a liquid moving in a
confined space under pressure) fluid to
Systems power hydraulic machinery
a. Integumentary – thin epidermis b. Skeletal –
mesodermal endoskeleton of calcareous plates • Medaporite = sieve plate connecting WVS to
→movable or fixed) c. Water vascular – derived outside seawater
from coelomic pouches • Stone canal = canal connected to
d. Nervous – a diffused net typically of three rings madreporite leading to the ring canal
center on mouth region with radiating →reinforced with spicules or plates of
branches; ectoneural system →receive sensory
CaCO3)
input from epidermis)
• Ring canal = canal forming a ring around the
e. Excretory – isotonic, some make use of
esophagus of most echinoderms
amoebocytes from Tiedemann bodies f. Respiratory –
skin gills, cloacal respiratory trees in Holothuroidea • Pollan vesicle = accessory fluid storage
g. Digestive – usually complete with anus on aboral structure, associated with ring canal
surface • Tiedemann’s bodies = filter fluid from WVS
h. Circulatory – water vascular system and the into main body cavity
coelom • Perivisceral coelom = main body cavity
i. Reproductive – asexual reproduction common; • Radial canal = canals radiating symmetrically
sexes usually separate. Fertilization is external.
from ring canal

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Zoology 111 – Invertebrate Zoology Jules Raposas
 - Ambulacral zones = regions penetrated by collagen synthesis,
tubular extensions of WVS transport of oxygen and nutritive materials,
- Ambulacral ossicles wound repair
• Ampullae = bulb-shaped structures connected
to the radial canals Echinodermata taxonomy
• Tube feet = thin walled tubular structures used
a) Class: Crinoidea – sea lilies and feather stars
for locomotion and feeding b) Class: Stelleroidea
1. Subclass: Ophiuroidea – brittle stars
• Ambulacral grooves →2000) 2. Subclass: Asteroidea – sea stars i.
Concentricyocloidea – sea daises
Function of the tube feet c) Class: Echinoidea – sea urhcins, heart urchins
1. Locomotion; attachment to substrate by and sand dollars
suction d) Class: Holothuroidea – sea cucumbers
2. Respiratory function
3. Primary site of secretion by simple diffusion Class Crinoidea
4. In some groups for chemoreception and food • Sea lilies and feather stars
collection • Arms branched; attached by stalk or free-
• Genes for vision in vertebrates • Suggests that moving
tube feet also function for light perception and • Mouth and anus on oral surface • No spines,
possible madreporite →attached to the ring canal), and
i. Inner surface is well ciliated, fluid in the
pedicellariae
WVS circulates
ii. Thin walled podia can function in gas - Pedicellaria = one of the minute pincer-like
exchange structures common to starfish and sea
urchins, used for cleaning and to capture
Hemal system tiny prey
• Appear like underwater exotic flowers
• Transports nutrients from coelomic fluid to
- Stalked crinoids →sea lilies)
gonads →asteroids and holothurians)
- Non-stalked comatulid crinoids →feather
• Excretions →asteroids and echinoids) • Axial
stars)
organ → spongy organic, major component of
hemal system; lies adjacent to stone canal;
Class Stelleroidea
produces coelomocytes • L: star
• Axial sinus → coelomic compartment • Star-like
housing the axial organ • Hemal rings → • Arms in multiples of 5; extends form central
vessels associated with the coelom and axial disc
gland • Lacks a stalk →not attached to a substrate)
• Perihemal canal → extend outward to the
gonads a) Subclass: Ophiuroidea
- Brittle stars and basket stars
• Coelomocytes → phagocytosis, pigment and

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- Arms distinct from central disc. locomotion)

Ambulacral grooves closed, tube feet - Pedicellariae and anus absent
without suckers →not used in - Autotomy: fragment and grows - has oral
 and aboral surface • Skeleton rigid →plates fused), mouth parts
present
b) Subclass: Asteroidea • Pedicellariae with three →3)-jaws •
- Sea stars and starfish
Spines are movable
- Arms are not distinct from central disc. • Ambulacral grooves closed
Ambulacral grooved open, tube free
• Has oral and aboral → contains the star
with suckers →used in locomotion) relief)
- Pedicellaria are present
- Oral has Aristotle structure
- Has oral and aboral surface

• Concentricycloidea Class Holothuroidea

- Now believed to be under the subclass • Sea cucumbers
asteroidea • Elongated body with no arms, spines, or
- Circular, flat, no arms, more resembles a pedicellariae
jellyfish medusa →with velum) or
• Skeleton only of microscopic plates • Mouth
flower rigged by retractile tentacles →modified tube
- Double-ringed WVS →classified it under feet
asteroidea)
• Pedicellariae absent, madreporite • True
- Body supported by series of overlapping evisceration
skeletal plates →ossicles)/calcareous
spines Developmental Stages
- Digestive system maybe absent; if present, 1. Doliolaria → adult sea lily (Crinoidea)
no anus 2. Bipinnaria → brachiolaria → sea star (Asteroidea)
- Xyloplax turnerae 3. Echinopluteus → sand dollar, sea urchin
(Echinoidea)
4. Ophiopluteus → brittle star (Ophiuroidea)
Class Echinoidea
5. Auricularia → sea cucumber (Holothuroidea)
• Sea urchins and sand dollars
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Aschelminthes → Cavity worms both ends

• Closely resembles the nematodes in Abundance
morphology • Most abundance multicellular animals
• Lost its taxonomic value for it does not come • 1-4M per m2in shallow water
from same ancestor as nematodes • Can be 90,000 in a rotten apple
Characteristics
• Vermiform Distribution
• Exhibits eutely →programmed to produce/ • Polar regions to tropics; mountaintops to seas
divide at a certain number) • Pseudocoelomic →no • Marine, freshwater
peritoneum) Groups • Bore holes of gold mines in South Africa
1. Gnathifera = jawed animals
2. Cycloneuralia = molting aschelminths Lifestyle
• Free-living or parasitic
Cycloneuralia →Gr: cyclo – circular; neura – - High densities near plant roots
nerve) - Species-specific relationships
• Circular braine or nerve ring around anterior - Economically important
digestive tracts • Eutely
• Exhibits eutely →1000 cells) - Cryptobiosis-ability to withstand periodic
desiccation
• Molting aschelminths
• Protonephridia as typical excretory organ Body wall
A) Cuticle
Ecdysozoa B) Epidermis
• Clade of organisms that molth C) Musculature
• Molting regulated by ecdysone
Cuticle
Nematoda • Thick, multi-layered covering
Defining characteristics • Non-cellular →made of collagen) •
• Roundworms →vermiform) Permeable to water and gases
• Amphids on the head-paired, lateral sense • Composed of network of inelastic fibers
organs - Can decrease/increase lattice angle
• Trellis-like arrangement permits bending,
General characteristics stretching and shortening • Functions:
• Roundworms
- Protection
• ~20,000 species
- Selectively permeable
• Ubiquitous - Covers body externally and internally
• Unsegmented - Molted/ shed
• Acoelomate and pseudocoelomate • 1-
- Includes several layers →Annuli &
2mm, no external segmentation • Tapered at
Furrow)

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• During development →has 4 larval stages), sheds and is re-secreted 4x during each
 development and in reproduction
• Unlike arthropods, nematodes continue to i. Feeding is by mouth pump sometimes
increase in size between molts assisted by
• Parasitic forms – animal is enclosed within 2 stylets while strong
envelopes before completing 2nd molt muscle closes the anus
- The outer envelope is called sheath ii. Movement is sinusoidal
- Exsheathment occurs only when eaten by iii. Copulation assisted by male
suitable host copulatory
accessories
Epidermis
Nervous system and sense organs
• Functions
1. Secretes cuticle • Nerve ring plus associated ganglia • Four →4)
2. Stores nutrients major longitudinal nerve cords
3. Bears fibers → tonofilaments – can be for - Dorsal, ventral, at least 1 pair of lateral
mechanoreceptors nerve cords
a) Brain
Musculature - Collar-like
• No circular muscles - circumpharyngeal
• Muscle layer is composed entirely of b) Nerve cords
longitudinal fibers - Suspending dorsal, ventral, lateral c)
• Body cavity is a hemocoel/pseudocoel Sensilia
- Small/non-existent in small, free living - Bear ciliated dendrite enclosed in
organisms specialized cuticularized part in the body
- Fluid in the cavity is pressurized and wall
functions as hydrostat d) Papillae
- Arranged in 3 rings around the mouth
• 2 factors contribute to maintain high internal - Mechanoreceptors
pressure e) Setae
1. Cuticle cannot expand to relieve stress - Elongate cuticular bristles on the head and
2. Muscle is always in partially contracted the body
state, trying to compress an - Touch-receptors
incompressible fluid f) Amphids
- Free-living aquatic and marine nematodes
Locomotion - Pouch or tube-like invaginations of the
• Sinusoidal movement: undulations are in the cuticle open to the exterior and
dorsoventral plane; eel-like but not side to containing ciliated receptor cells
side; no antagonistic actions of a circular - Paired and found on the head
muscle so they push against their cuticle - Mechano and chemoreceptors
• Related to cuticle and fluid pressure - Animal g) Phasmids
encounter problems in feeding, locomotion,

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- Found in the tail regions of some nematodes function is uncertain

- Paired unicellular glands
- Chemosensory, secretory, excretory; best Feeding habitat and digestive system •
developed in parasites Carnivores and herbivores
h) Ocelli • Deposit-feeding
- Laterally, on the sides of the pharynx; - Turbatrix aceti – lives in the sediment of
 non-pasturized vinegar - Receives digestive wastes and sperm
• Digestive tract - Evaginated to form 2 pouches
1. Foregut – consists of the buccal cavity - Short; shaped like a pointed curved blade
2. Midgut – consists of the intestine 3. - Female vagina opens to gonopore
Hindgut – rectum
• Mouth is at anterior end Unusual reproductive characteristics 1)
• Teeth and stylet may be present; especially in Sperm
carnivores - Amoeboid rather than flagellated 2)
• Mononchus papillatus Chromosomal diminution
- Toothed terrestrial nematode - Specific regions of the DNA are destroyed
- Attaches lips to prey, puncture the skin and - Somatic cells lose some genetic material
sucks out the content of the prey through - Cells divide parallel to the cleavage plane
muscular pharynx i. Germ cell undergo meiosis
- Consumes as many as 1000 other ii. Somatic cells have degenerated
nematodes in a span 18 weeks chromatin
→disintegrated upon
Anatomy of digestive and reproductive systems division)

Excretion Development
• Simple diffusion across the body wall • • Exhibit eutely
Ventral cell • Cleavage is determinate →mosaic) - Cell fates
- Excretory gland called Renette cell are permanently set at first cleavage
- Protrudes into the hemocoel • Life cycle
- Has a neck-like duct that opens into a - Egg → 3 juvenile instars → adult i. Instar –
midventral pore stage in the life cycle between successive
Reproduction and development molts; also used
• Gonochoristic; hermaphroditic; arthropods
parthenogenetic ii. Larvae emerge from sturdy eggs as
• Females produce pheromones miniature
• Fertilization is internal, with copulation adults
• Male rectum is a multipurpose cloaca

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iii. Some may enter a stage of Taxonomy

developmental arrest Class Aphasmida/Adenophorea
• Dauer larva: state of developmental arrest
- Essentially inactive No phasmids (caudal sensory organs);
- Has a very low metabolic rate, and cannot amphids much modified externally except in
feed parasitic forms; amphids are chemosensory receptors
for female pheromones. Excretory system
Embryo → L1 → may proceed to L2 or become rudimentary or absent; coelomocytes and mesenteric
Dauer) tissue well developed.

Order Trichurida/Trichocephalida
Class: Adenophorea →Aphasmida Order: • Esophagus a very long, fine tube embedded for
Trichocephalida →Trichurida Family: most of its length in a column of glandular cells
Trichuridae (stichosome) and short muscular region; females
with one ovary; males with one spicule or none.
Trichinella spiralis Capillaria
• Trichina worms. The cysts are at first very delicate • These are very fine filamentous worms, between 1
but gradually thicken and become lemon-shaped and and 5 cm long. The narrow stichosome esophagus
are located parallel to the muscle fibers. As a rule, occupies half of the body length. In the ova, bipolar
only 1 or 2 worms are encysted in a cyst but as plugs can be seen at the polar ends. Hosts of
many as 7 have been seen. After 7 or 8 months, the Capillaria include avians and mammals. Life cycle
cyst walls start to calcify, beginning at the poles. similar to Trichiuris.
After 18 months, the entire cyst becomes calcified
and appears as a hard calcareous nodule. The Class Phasmida/Secernentea
enclosed worm usually degenerates and dies after
some months. After encysting in the flesh, no further Phasmids present; amphids simple pores;
development takes place until a excretory system present or rudimentary;
susceptible animal eats the flesh, whereupon the coelomocytes (6 or less) and mesenteric tissue
worm matures and begins reproducing in a few days. weakly developed.

Trichiuris trichiura Order Ascaridida

• Whip worms. Parasitic worms found in the • Esophagus bulbed or cylindrical; vagina elongate;
intestines of man and other mammals. They are mouth usually with 3 or 5 lips; males usually with 2
named such because their body resembles a whip, spicules; tails of male not spirally coiled but usually
particularly the very long stichosome esophagus. The curled ventrally; no true bursa, but alae or
mouth is found at the tip of the esophagus. longitudinal ridges may be present.
Infestation in man, especially children, takes place
through ingestion of infected soils. The worm injects Superfamily Ascaridoidea
a fluid on the host tissue and subsequently takes in • Cervical papillae present; mostly large, stout
the dissolved tissue. The eggs possess bipolar plugs. polymyarian worms; males with 2 spicules; tail
curled ventrally; with or

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without lateral alae; esophagus muscular with or the length of the anterior end. In mature female, the
without a bulb. level where this pore is situated is distinctly
constricted.
Ascaris lumbricoides • X-section: The x-section of Ascaris shows the body
• Large opaque worm, parasitic in the intestines of wall consisting of three layers: (1) the cuticle, the
some mammals, including man. Mature specimens non-living transparent outermost layer; (2) the
from hogs are indistinguishable morphologically hypodermis consisting of scattered nuclei and fibers
from those of man but are physiologically different. but without cell membranes (syncytium); and (3) the
• External features. The male is generally smaller thick muscular layer composed of prominent muscle
than the female; its posterior end is curved and, in cells whose internal extensions appear as thin-walled
some specimens, the 2 pineal setae may be sticking sacs or bulbs into the pseudocoel. Because of the
out of the anus. The mouth is at the opposite end, presence of the 4 lines on the body wall, the
surrounded with 3 lips. Two very distinct lateral muscular layer is divisible into 4 quadrants. The
lines are found throughout the length of the body. dorsal and ventral lines contain longitudinal nerves
Two other lines, the mid dorsal and mid-ventral lines, while the lateral lines have the excretory ducts.
are less distinct. • The pseudocoel is really the confluent vacuoles of
• The female also has the features of the male except giant cells, not a single cavity. In the x-section, the
for the absence of the curvature of the posterior end protoplasmic strands separating the vacuoles may
of the pineal setae. The female genital pore (or appear as faint
vulva) is situated on the ventral line, about a third of lines criss-crossing the space known as the
 pseudocoel. Extending along the body length are 2 prominent
• The section of the intestine is a flattened hollow lateral lines and less distinct dorsal and ventral lines.
structure lined with a single layer of columnar cells. • The female, with a straight posterior end is
In the male, the large seminal vesicle containing generally larger than the male. Locate the mouth,
spermatozoa and several sections of the coiled and the lateral, dorsal and ventral lines. On the
deferent duct and testis may be recognized. In the ventral line, at about the anterior third of the body,
female, 2 large cavities containing eggs are the uteri. look for a very small opening, the genital pore.
The oviducts and ovaries are represented by several • Internal Anatomy
circular sections consisting of the triangular cells a. Male
meeting at a common center, which is clearly hollow • The digestive tube is a straight tube extending from
in the oviduct. the mouth to the anus. Immediately, posterior to the
mouth is the short, muscular esophagus that leads
Ascaris suum into the long intestine. Posteriorly, the intestine
• Pig ascaris. narrows into a short cloaca, which leads into the
• External Anatomy. The male ascaris is generally anus. The reproductive system consists of a single
smaller than the female, and provided with a curved tube. The tube ventral to the intestine at the posterior
posterior end. On the anterior end, look for the end of the body is the seminal vesicle. Follow the
mouth. Near the posterior end is the cloacal opening seminal vesicle posteriorly and observe that it
where 2 small rods, the pineal setae protrude. narrows into the ejaculatory duct, which leads into
the cloaca. Anteriorly, the seminal vesicle

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narrows into the coiled vas deferens. Carefully lift tail of female usually slender and pointed
the vas deferens and note that it is continuous with a
finer tube, the testis. Enterobius vermicularis
• Pinworms. Lives in the caecum, appendix and parts
b. Female of the intestine from which the female migrates to
• The female digestive tube has no cloaca, thus the the rectum or the perianal region to deposit the eggs.
intestine opens directly into the anus. The This activity in the rectum of the host body causes
reproductive system is an inverted Y-shaped tube. severe itching. The eggs are clear and unstained,
Connected to it is a short tube, the vagina, which is about 55 x 60 micra and are flattened on one side;
formed by the union of the left and right uteri which airborne or conveyed by the hands.
courses posteriorly and coils back anteriorly as the • Through the adult’s cuticle, which is
fine oviduct. It ends into a very fine thread like tube, semitransparent, one sees the esophagus with the
the ovary. bulb at its posterior end and the uteri and
coiled ovaries. The head is set off by lateral
Toxocara canis expansions of the cuticle called cephalic alae and has
• Common ascaris of dogs being especially prevalent 3 small lips. The female tapers at both ends, tail is
in puppies. Dogs become infected by swallowing the drawn out into a long fine point hence the name
eggs. The larvae migrate to the body as do those of pinworm.
Ascaris lumbricoides in people. Dogs acquire an • The male is minute and the tail is curled with
immunity as a result of the infection, and after 3 to 4 bursa-like expansion with one spicule.
months the worms are cast out and susceptibility to
further infection is lost. Heterakis gallinae
• Phasmid worms that live in birds, particularly in
Superfamily Oxyuroidea the cecum of barnyard and wild fowls. Eggs, which
• Cervical papillae absent; mostly small or of are passed in the feces of infected birds, are
medium sized transparent meromyarian worms; swallowed and the young that hatch from them in the
males with one or two spicules; esophagus bulbed; small intestine move on to the ceca. They do not
 seriously injure the fowls but are of great economic countries. The larvae of this species are usually ¼
importance since they carry with them the protozoan mm long. During the daytime they live in the lungs
parasite Histomonas meleagridis, which is the and larger arteries, but at night they migrate to the
causative agent of the disease in turkeys known as blood vessels of the skin. Mosquitoes that are active
blackhead. at night suck up these larvae with the blood of the
infected person. The larvae develop in the
Superfamily Filarioidea mosquito’s body, becoming about 1 mm long, make
• Parasites of the body fluids and tissues of their way into the mouth parts of the insect, and enter
vertebrates; transmitted by arthropods; vulva of the blood of the mosquito’s next victim. From the
female near the anterior end; slender worms. blood they enter the lymphatics and may cause
serious disturbances, probably obstructing the lymph
Wuchereria bancrofti passages. This results in the disease called
• Parasite in humans and widely spread in tropical elephantiasis. The limbs or other regions of the body
swell up to an

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enormous size, but there is little pain. However, the end while the shorter male bears posteriorly the
disease can be fatal. large, ventrally directed bursa copulatrix for holding
the female during copulation.
Dirofilaria immitis • In man, infecting hookworms are called
• Dog heartworm. They are found as tangled masses Ancylostoma duodenale. These are rather stocky
in the right ventricle of the dog’s heart. They worms about an inch long with well developed
occasionally infect humans. bursa, very long needle like spicules and a
conspicuous goblet or cup
Order Strongylida shaped buccal capsule guarded ventrally by a pair of
• Usually meromyarian; males with 2 spicules and chitinous plates which either bear teeth as in
with a true bursa supported by 6 paired rays and one ancylostomes or have a blade-like edge as in
dorsal ray which may be divided; mouth simple, necators. Ancylostoma duodenale
without lips or with a buccal capsule; esophagus has two well-developed teeth on each plate with a
muscular, club rudimentary third one near the median plane; head is
shaped, or cylindrical; eggs thin-shelled and coarse and only slightly bent dorsally.
colorless.
Necator
Ancylostoma • Necator americanus is the most important species
• Hookworms. These nematodes live in the intestines throughout the tropics despite the scientific name,
of man and some other mammals. The body is which can be
encased in a cuticle with its anterior and curved confused to have originated from America but it is
dorsally. The longer female has a pointed posterior not.
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