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Title: 1st: Introduction to Invertebrate Zoology
Description: 1st year Introduction to Invertebrate Zoology notes, University of Exeter
Description: 1st year Introduction to Invertebrate Zoology notes, University of Exeter
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1: INTRODUCTION
2
2: NATURAL SELECTION
2
2: CLASSIFICATION
3
2: PHYLOGENIES AND LINKING PHYLOGENY WITH CLASSIFICATION
3
2: THREE DOMAINS
5
3: PROKARYOTES
5
3: EUKARYOTES
7
3: SERIAL ENDOSYMBIONT THEORY (SET)
7
3: PROTISTS
8
4: ANIMAL DIVERSITY
9
18: THE GREAT ANIMAL RADIATION
11
5: PORIFERA
14
5: CTENOPHORA
14
5: CNIDARIA
15
5: ACOELA
16
6: LOPHOTROCHOZOA
16
6: PLATYHELMINTHES
16
7: ROTIFERA
18
7: ECTOPROCTA
19
7: BRACHIOPODA
20
8&9: MOLLUSCA
20
10: ANNELIDA
28
12: ECHINODERMATA
31
13,15,16,17,18,&19: ARTHROPODA
35
17: NEMATODA
52
Joanna Griffith (2017)
1: INTRODUCTION
Why study invertebrates?
○ There are 5 million different species, and only 5% of them are vertebrates
○ Invertebrates are more diverse than vertebrates in form, ecology, and
morphological function
● Zoology: the study of animals
○ Animals are usually multicellular, heterotrophic, and eukaryotic, with tissues
that develop from embryonic layers, and they do not have cells with
supporting cell walls
■ Definition is not straightforward, there are always exceptions
○ The form of an animal can relate to:
■ Type of environment
■ Size of animal
■ Mode of existence
● Adaptation: animal form and function is correlated with the environment at all levels
of organisation
○ Convergent evolution: different evolutionary history, but similar solution
developed for the same problem (eg
...
Panthera leo)
○ Genus is always capitalised, whole binomial name in italics or underlined if
handwritten
○ Name tends to be informative about the distinguishing features of the
organism
● Hierarchical classifications
○ Domain
■ Kingdom
● Phylum
○ Class
■ Order
● Family
○ Genus
■ Species
● Taxon: taxonomic unit at each level
● Classification is informed by phylogenetics, but they are not the same thing
Cladistics
● Classification based on phylogeny (the evolutionary development of a species)
● Clades contain ancestral species and all of their descendants
○ Helps to identify groups that share a unique common ancestor
○ Nestled within larger clades
● A taxon from the Linnaean system is only equivalent to a clade if the clade is
monophyletic (a group of organisms all descended from a common ancestor)
● Monophyletic group: an ancestral species and all of its descendants
● Paraphyletic group: an ancestral species and some of its descendants
● Polyphyletic group: taxa with different ancestors
● Clades are inferred from shared derived characters (characteristics that differentiate
them from other clades, such as hair in mammals)
○ Not to be confused with shared ancestral characters (characteristics passed
on by the common ancestor, such as a backbone)
■ However, these shared ancestral characters can be used to
differentiate between larger clades (eg
...
wings in
bats and birds are homologous as flight structures)
■ Synapomorphies: characteristics present in an ancestral species and
shared exclusively (in more or less modified forms) by its evolutionary
descendants
○ Analogous structures: features in different organisms that appear similar but
are actually the result of convergent evolution (eg
...
brachiopods or
ectoprocts) or have a distinctive development stage as
trochophore larvae (eg
...
thermophiles, halophiles
○ Eukarya
■ All organisms with eukaryotic cells
● Cells with a nucleus
● Unicellular organisms, multicellular organisms, plants, animals,
fungi
○ Reflects that life is dominated by single-celled organisms
○ Kingdoms of Protista and Monera are no longer recognised
--------------------------------------------------------------------------------------------------------------------------●
3: PROKARYOTES
●
Collective biomass is 10x that of eukaryotes
Joanna Griffith (2017)
●
●
●
●
●
●
●
●
●
Huge genetic diversity
Most are unicellular, some are colonial
Lack membrane-bound organelles
Range of nutritional modes
○ Autotrophs
■ Photoautotrophs
● Light as an energy source, CO2 as a carbon source
● Eg
...
certain prokaryotes such as Sulfolobus
■ Heterotrophs
● Photoheterotrophs
○ Light as an energy source, organic compounds as a
carbon source
○ Eg
...
many prokaryotes such as Clostridium
Three main shapes
○ Rods (bacilli), eg
...
Staphylococcus aureus
○ Spirals (spirilla), eg
...
Giardia), Parabasalids (eg
...
Trypanosoma)
○ Chromalveolata
■ Based on DNA sequencing
■ Originated a billion years ago
■ Split into Alveolates and Stramenopiles
○ Rhizaria
○ Archaeplastida
■ Based on molecular systematics and cell structure
■ Eg
...
surface of embryo, outer layer of the animal, the central nervous
systems of some phyla
○ Endoderm
■ Eg
...
jellies
○ Triploblastic
■ Has three germ layers
■ Eg
...
earthworms
■ Pseudocoelomates
● Formed from mesoderm and ectoderm
● Eg
...
flatworms
Development
○ Protostome
■ Eg
...
echinoderms and chordates
■ Radial and indeterminate cleavage in the eight-cell stage (removing a
single cell has no effect on the finished organism and can even lead to
several blastopores developing)
■ Folds of archenteron form the coelom
Joanna Griffith (2017)
■ The blastopore develops into the anus, the mouth is secondary
Why is there so much functional diversity?
● Regulatory genes contain sets of DNA sequences called homeoboxes, which have
been very well conserved throughout evolution, and similar versions are present in all
animals
○ Despite this, the animal kingdom is very diverse
● mass/surface area ratios
○ V=pir2h
■ Assuming constant shape, volume is proportional to length3, and
surface area is proportional to length2
■ Need for oxygen is proportional to volume, which is proportional to
length3
■ External surface area available to absorb oxygen is proportional to
length2
○ Surface area must keep pace with volume
■ Larger animals must be more morphologically complex in order to
absorb enough oxygen
● Tensile strength
○ Mass = load, which is proportional to length3
○ Tensile strength is proportional to cross-sectional area, which is proportional
to length2
● Compound eyes
○ Not primitive, just suited to smaller animals
■ Reduces refraction of light by providing a larger aperture
● Scale laws
○ Size affects speed/ability to jump/ability to fly/etc
...
5bya
● Tappania (eukaryote - protist) 1
...
6bya
● Marella (eukaryote - animalia), 0
...
Charniodiscus and
the sea pen), some don’t
○ Probably surface grazers
■ Bottom of ocean thought to be a microbial mat on top of anoxic
sediments
■ Trace fossils from 550 mya show no sign of burrowing
● The Cambrian explosion (Paleozoic era, 542-251 mya)
○ Rapid diversification between 535 and 525 mya
○ 50% of all extant phyla emerged in this period
○ Soft-bodied fossils found in the Burgess Shale (eg
...
7 - 2
...
Ediacaran fossils from Dengying formation (555 mya)
● Inferred increase in O2 accompanies the evolution of complex life
○ Aerobic respiration (36 ATP molecules per glucose) is more productive than
anaerobic respiration (2 ATP molecules per glucose) and is now easier
○ More oxygen = more energetic lifestyles
■ More predation, larger organisms
● Evidence for increased oxygen
○ Increase in carbon burial
○ The solubility of rare minerals changes in the presence of oxygen (eg
...
shells, hard exoskeletons
(eg
...
Treptichnus pedum)
○ Oxygen produces ozone, which reduces the incidence of UV in surface
waters and makes colonisation of land possible
The snowball Earth hypothesis
● Growing polar ice caps cause the Earth to freeze over, volcanic outgassing leads to a
build-up of volcanic CO2 which melts the ice, high phosphorus levels in oceans cause
a huge global algal bloom
○ Photosynthesis of blue-green algae = O2 levels increase
● Positive feedback (cooling)
○ Ice formation at poles increases the albedo effect, greater heat reflection,
runaway cooling exacerbated by having land masses at the equator,
weathering keeps removing CO2 from the atmosphere even with icy poles
● Negative feedback (warming)
○ Volcanoes vent CO2, no removal by rain
○ When icy, no rock weathering to bind CO2 to silica
○ Melting predicted to be rapid
The fossil record
● Many phyla appear suddenly as well-differentiated classes, eg
...
scorpions)
○ Paleozoic era finishes with the Permian mass extinction event, ‘the great
dying’
● The Mesozoic era (251-65
...
5-present)
○ Mass extinction event at Cretaceous/Tertiary boundary
■ Dinosaurs disappeared, frogs survived
○ Rise of the mammals, exploited vacant niches
--------------------------------------------------------------------------------------------------------------------------5: PORIFERA
● About 5000 species of sponges
● marine/freshwater
● Large size range, from mm to m
Body plan
● Diploblastic (only two embryonic cell layers)
● Radially symmetrical or asymmetrical
● Lack true tissues, unlike other animals
○ True tissues: groups of similar cells that act as functional units and are
isolated from other groups of cells by membranous layers
○ Allows for a process called reaggregation, where the sponge can reform even
after having been destroyed (eg
...
sea gooseberry (Pleurobrachia), warty sea walnut (Mnemiopsis leidyi),
venus girdle (Cestum veneris)
About 100 species, all marine
Joanna Griffith (2017)
Body plan
● Radially symmetrical
● Diploblastic
● Ciliated longitudinal rows used for propulsion
● Long, branched, contractile tentacles
○ Have sticky cells called colloblasts
■ Used to trap prey
● Many species are bioluminescent
Reproduction/development
● All species are hermaphroditic
--------------------------------------------------------------------------------------------------------------------------5: CNIDARIA
● Sister group to Eumetazoa
● Wide range of motile and sessile forms
● About 10,000 species, including corals, anemones and jellies
Body plan
● Has true tissues
● Diploblastic
● Radially symmetrical
● Gastrovascular cavity is also known as a coelenteron
Nutrition/digestion
● Carnivorous, predatory lifestyle
● Extensible tentacles capture prey and pass it into the gastrovascular cavity
○ Some Cnidarians can also perform extracellular digestion
● Tentacles may be armed with cnidocytes (stinging cells unique to Cnidaria)
○ Cnidae are capsule-like organelles
■ Nematocysts are specialised cnidae with stinging threads that
penetrate the body of prey
■ Other types of cnidae produce sticky/tangling threads
Nervous system/sense organs
● Muscle and nerve net
○ Bundles of microfilaments are arranged into contractile fibres in the epidermis
and gastrodermis
● No brain, simple reflex pathways
● Nerve cells interact with sensory and contractile cells to coordinate movement
Hydrozoa
● Commonly colonial organisms
○ Multiple individuals combine to make “one” organism
● Have both polyp and medusa stages, and normally alternate between them
Scyphozoa
● “True jellies”, sea nettles
● All marine
● Mostly larger jellies
○ Many are bioluminescent
Joanna Griffith (2017)
● Predominant medusa stage
Cubozoa
● Box jellies and sea wasps
○ Box-shaped medusa stage
● All marine, in the topics
● Strong swimmers
● Extremely potent venom
● Active hunters, prey mainly on fish
● Complex eyes along fringe of tentacles
Anthozoa
● “Flower animals”, such as sea anemones, most corals, sea fans
● Only occur as polyps
Myxozoa
● Parasitic class
● Unicellular, “protist”
● Highly reduced genome
● Has a polar capsule that is very similar to nematocysts
○ Evolution can simplify animal organisation
--------------------------------------------------------------------------------------------------------------------------5: ACOELA
--------------------------------------------------------------------------------------------------------------------------6: LOPHOTROCHOZOA
Bilateral symmetry
Triploblastic
Clade identified by molecular data
○ Wide range of animal body forms
■ 18 phyla
■ No unique morphological features shared by all members
● Named after lophophores, crowns of ciliated tentacles, found in some members of
the clade, and trochophore larvae, which as small, translucent, spinning-top-shaped
larvae with a prominent circle of cilia, also found in some members of the clade
○ Some members of the clade lack both of these features
--------------------------------------------------------------------------------------------------------------------------●
●
●
6: PLATYHELMINTHES
●
●
●
Flatworms, flukes, and tapeworms
About 20,000 species
Consist of two lineages
○ Catenulida
■ Chain worms
■ Clade of 100 species
○ Rhabditophora
Joanna Griffith (2017)
■
■
■
■
Clade of about 20,000 species
Includes free-living and parasitic species
Both marine and freshwater species
Eg
...
schistosoma
○ Causes schistosomiasis (200 million people infected)
○ Live in blood vessels for about 40 years
○ Reproduce sexually in a human host, eggs exit the host in faeces and
develop into ciliated larvae in an intermediate snail host, undergo asexual
reproduction and become motile larvae that can penetrate human skin
Cestoda
● Tapeworms
● About 4000 species
● All species are parasitic
○ Normally infect two host species in their lifecycles, mainly infect vertebrates
● No gastrovascular cavity, as they absorb nutrients through their skin
● Have a scolex with hooks and suckers
● Have proglottids with reproductive units
● Common cestodes in humans:
○ Beef tapeworm (Taenia saginata)
○ Pork tapeworm (Taenia solium)
○ Fish tapeworm (Diphyllobothrium)
○ Dog tapeworm (Dipylidium canium)
○ Dwarf tapeworm (Hymenolepis nana)
--------------------------------------------------------------------------------------------------------------------------7: ROTIFERA
● Multicellular, have a fixed number of highly differentiated cells
Body plan
● Pseudocoelomates
● Have a crown of cilia (a characteristic that allows them to be classified as
Lophotrochozoans)
Joanna Griffith (2017)
● Specialised organ systems
● Trunk contains visceral organs
● Cloacal bladder collects excretory/digestive waste
Nutrition/digestion
● Alimentary canal with two openings
● Pharynx (called a mastax) with intricate jaws
● The corona channels water currents into the mouth, used for feeding (and
locomotion)
● Rotifers rotate around their segmented, telescopic foot (1-4 toes) to feed
Respiration/circulatory system/excretion
● Protonephridia (flame bulbs) used for osmoregulation and excretion
Reproduction/development
● Life cycles of the three Rotifera classes
○ Gonochoristic (separate male and female sexes) reproduction in Seisonidea
○ Parthenogenesis (reproduction from an ovum without fertilisation) alternating
with sexual reproduction in Monogononta
○ Obligate parthenogenesis in Bdelloidea
■ Exclusively asexual reproduction
● Nearly all other organisms have a sexual phase in their life
cycle
○ Asexual lineages often arise, but they are normally
short-lived due to a slower rate of evolution compared
to parasites and an accumulation of deleterious
mutations
○ Interactions with parasites are thought to provide
selective pressure to maintain sex
○ Bdelloidea may escape fungal pathogens by using a
dessicated resting stage (anhydrobiosis) and dispersal
--------------------------------------------------------------------------------------------------------------------------7: ECTOPROCTA
“Moss animals”
About 4000 species
Small (about 0
...
the trembling sea mat, Victorella pavida
○ Only found in Swanpool, Falmouth
Body plan
● Lophophore (crown of ciliated tentacles)
● Coelomates
●
●
●
●
●
●
●
Joanna Griffith (2017)
Having a coelom is advantageous as it provides a hydrostatic skeleton and
allows for more efficient circulation of nutrients
○ U-shaped alimentary canal
○ No head
○ Hard exoskeleton
--------------------------------------------------------------------------------------------------------------------------○
7: BRACHIOPODA
About 350 species
All marine
Sessile bottom dwellers
Eg
...
clams, oysters, squids, octopuses, snails
● One of the largest phyla (about 93,000 species)
● Huge size range, from microscopic to 18m
● Marine, freshwater, terrestrial
Body plan
● Despite external differences, they are all built with the same body plan
○ All are soft-bodied (no skeleton) and most secrete a hard shell
● Coelomates
● Mantle
○ The dorsal surface is covered by a shell that protects the internal organs
○ The mantle (epidermis) secretes the shell
●
Joanna Griffith (2017)
Epidermal secretions are continuous so the shell increases in size and
repairs itself throughout the mollusc’s life
● Retractor muscles
○ Retractor muscles are organised in pairs, and each pair is attached to the
inner surface of the shell and foot
● Mantle cavity
○ The shell and underlying mantle overhang the body, creating the mantle
cavity
○ Contains a pair of gills and a pair of nephridia
Nutrition/digestion
● Grazers, carnivores, and filter feeders
● The mouth contains a feeding organ called a radula, which allows most molluscs to
graze
Respiration/circulatory system/excretion
● Gills
○ Water enters the lower mantle cavity, passes through the gills and back,
pushed by the beating of cilia
■ Two blood vessels run over the gills and are oxygenated
● The heart
○ Open circulatory system (except in Cephalopods)
■ The coelomic cavity surrounds the heart
○ The heart consists of a pair of posterior auricles and a single anterior ventricle
● Excretory organs
○ Pair of tubular metanephridia (kidneys)
■ One end connected to the pericardial cavity, the other connected to a
nephridium
● The pericardial cavity receives waste from the heart, waste
then passes into the nephridium and urine exits through the
mantle cavity
Nervous system/sense organs
● A nerve ring loop around the oesophagus
● Pairs of nerve cords extend ventrally and dorsally
○ Ventral pair innervates the muscles of the foot
○ Dorsal pair innervates the mantle and visceral organs
● Sense organs include tentacles, eyes, touch receptors, and chemoreceptors
Reproduction/development
● Most are gonochoristic, some are hermaphroditic
● Gonads present on either side of the coelom
○ ‘Ripe’ eggs and sperm break into the coelomic cavity and are transported to
the external environment through the nephridia
■ Fertilisation occurs in seawater
● Most marine and aquatic molluscs have planktonic trochophore and veliger larval
stages
Locomotion
● The epidermis is covered by cilia and mucous gland cells
○ Mucus lubricates the substratum under the foot for locomotion
■
Joanna Griffith (2017)
Formation of pearls
● Pearls form inside the shells of certain molluscs as a defense mechanism against a
potentially threatening irritant (eg
...
snails, slugs, limpets, whelks, conch, sea hares, periwinkles, sea butterflies
● 75% of all mollusc species are gastropods
Joanna Griffith (2017)
●
●
●
●
The only molluscs to have colonised land
Body plan
○ Shell morphology
■ Have a heavy, univalve, coiled shell (coils may reduce or disappear in
adult stages)
■ Planispiral to asymmetrical shell coiling
● Ancestral planispiral shells were huge and heavy, leading to
the development of asymmetrical coiling
■ Shell consists of four layers
● Outer layer is conchiolin or conchin
● Inner layers consist of calcium carbonate
■ Shell colours result from pigment in the calcareous layers of the shell
■ Retractor muscles close the shell and the operculum acts as a door or
lid
○ The anus and the nephridiopore open near the pneumostome (lung opening)
○ Torsion
■ Process of moving the mantle cavity to the front of the body
■ Pre-torsion: anterior head and posterior anus
■ During torsion (during the veliger larval stage), the visceral mass
twists 180°
■ Only occurs in gastropods
■ Post-torsion: left gill, kidney, and heart auricle are all on the right-hand
side and vice-versa
● Modern gastropods have lost their right gill, kidney, and heart
auricle
● Mantle cavity and anus open above the head and mouth
■ Advantages (heavily debated)
● The head can enter the shell first, then the foot with the
operculum to close the shell
● Allows entry of clean water onto gills
● Inhalant current can be sampled by osphradia sensory organs
in the mantle
■ Some nudibranchs undergo 90° detorsion as adults, likely for
sanitation issues
nutrition/digestion
○ Highly varied diets
○ Most are herbivorous (grazers/browsers)
○ Ciliary feeders (eg
...
the textile cone shell (Conus textile) uses its radula as a
venomous harpoon
respiration/circulatory system/excretion
○ Respiration via gills/skin
■ Terrestrial snails and slugs have a pallial lung
○ Well-developed circulatory system
Joanna Griffith (2017)
Nervous system/sense organs
○ Well-developed nervous system
● reproduction/development
○ Gonochoristic and hermaphroditic species
■ Elaborate courtship rituals, eg
...
copulation of shelled Helicidea
■ Vagina contains an oval dart sac which secretes a calcareous spicule
● When the snails are intertwined, one pushes the dart into the
body wall of the other
● Subclasses
○ Prosobranchia
■ Eg
...
sea hares (vestigial shell), sea slugs, nudibranchs (have cerata),
canoe shells
○ Pulmonata
■ Eg
...
mussels, clams, scallops, oysters
● Most marine, a few brackish water and freshwater
● Diverse size range, from about 2mm to 1
...
by razor clams to burrow
■ Foot can be inflated to anchor the bivalve into sediment
○ Eg
...
octopus, squid, cuttlefish, nautilus
○
Joanna Griffith (2017)
●
●
●
●
●
All marine, found in all oceans
Normally 2-70cm, but the giant squid (Architeuthis dux) can grow to up to 18m
○ The colossal squid (Mesonychoteuthis hamiltoni) is the heaviest known
invertebrate, at 500kg
Ammonites were dominant in the Mesozoic era until their Cretaceous extinction
Body plan
○ Tentacles
■ Nautilus have 38 tentacles, no suckers
■ Squid and cuttlefish have 4 pairs of arms and 1 pair of tentacles
● The inner arm is flattened and covered with stalked suckers
that are horny and toothed
■ Octopods have 8 arms and suckers
● Self-recognition mechanism between skin and suckers
prevents octopus arms from interfering with each other
○ “Spaghetti holding” behaviour
○ Form and function of the nautilus’ shell
■ All cephalopods have undergone a progressive loss of shell, except
nautilus, which still have an external shell
■ Nautilus shells are divided by transverse septa into internal chambers
● The septum is perforated in the middle by a cord of tissue
called the siphuncle
○ The siphuncle secretes gas into the empty chambers
and makes the shell buoyant
■ Limitations
● The nautilus can only reach certain depths before pressure
causes the shell to implode
● Not aquadynamic, creates drag through the water
■ The ancestor of all cephalopods had a shell
● Cuttlefish and squid have internal shells
● Octopods have no shells
● Nautiluses are the only cephalopods that have maintained an
external shell
○ Colour change
■ Chromatophores are present in the integument (protective outer layer)
● Small muscles are attached to the edges of the
chromatophores
● Different cephalopod species have chromatophores of different
colours
● The effect is enhanced by iridocytes or reflector cells
■ Some colour changes are associated with behaviour
● Camouflage
● Squid and octopods may darken when they are alarmed
● Colour displays associated with courtship
nutrition/digestion
○ All cephalopods are active predators, hunt a wide range of prey
○ Prey are captured using tentacles or arms
Joanna Griffith (2017)
Radula is present, but the jaws in the buccal cavity are more important
■ Tissue is pulled into the buccal cavity by the radula, and a pair of
salivary glands empty into the cavity
● Octopods also secretes poison and enzymes into prey
○ Cephalopod diets depend on their habitats
■ Squid feed on fish, crustaceans, and other squid
■ Cuttlefish swim along the bottom and feed on invertebrates such as
shrimp and crabs
■ Octopods tend to live in dens, so they can search for food or lie in wait
● They can capture clams, snails, crustaceans, and fish, and
paralyze them
respiration/circulatory system/excretion
○ Cephalopods are the only molluscs with a closed circulatory system
■ Blood remains separate from the fluid in the body cavity
Nervous system/sense organs
○ In cephalopods, the ganglia are concentrated and fused to form the brain
■ A cartilaginous cranium protects the brain
■ Separate regions of the brain control different areas
● Eg
...
Lumbricus terrestris (common earthworm)
○ Eg
...
Arenicola marina (lugworm)
○ Eg
...
Spirobranchus giganteus (Christmas tree worm)
○ Eg
...
Florida mudflats
○ Re-emergence in medicinal use
● Traditionally divided into 7 classes and subclasses
○ Phylogenetic analysis indicates most of these are either paraphyletic or
polyphyletic
Body plan
● Metameric (segmented)
○ Metameric structures confined to the trunk
○ The prostomium (head), peristomium (behind prostomium) and pygidium
(terminal part) are not segments
○ The formation of new segments takes place just before the pygidium
● Coelomates
○ The coelomic fluid functions as a hydrostatic skeleton
Locomotion
● Movement by peristalsis
○ Two sets of muscles, longitudinal and circular
○ Widening and elongation can be restricted to certain segments
○ Waves of peristaltic contraction pass down the length of the body and cause
elongation and shortening in each segment
○ Chitinous, paired, lateral bristles (chaetae) increase traction with the substrate
Polychaetes
● Largest class, about 10,000 species
● Most range from 5-10cm
● Mostly marine
● Wide range of lifestyles
○ Tube dwelling, burrowing, pelagic
● Body plan
○ Segmented worms with “legs” (chaetae/parapodia)
●
Joanna Griffith (2017)
●
●
●
■ Have many chaetae
○ Prostomium
■ Well differentiated
■ Specialised sense organs
■ Chitinous jaws (light and strong)
■ Bears eyes, antennae, and palps
■ Sometimes retractile
○ Peristomium
■ Surrounds mouth
○ Pygidium
■ Carries the anus
○ Have characteristic parapodia topped with chaetae
■ Paired, fleshy appendages that extend from the body segments
■ Sweep ground/water for locomotion
■ Can be converted into respiratory organs in larger worms
Nervous system/sense organs
○ Brain
■ Lies in the prostomium
■ Supplies nerves to the palps, antennae and eyes
■ The ventral nerve cord is doubled through the segments, although the
two cords are often fused
■ Presence of giant axons (1
...
in peacock
worms
● Increases reaction time by 40x
○ Retinal cup eye
■ Detects light intensity and source
○ Nuchal organs on head region
■ Detect food
○ Statocytes
■ Found in burrowers
■ Detect changes in orientation
Reproduction
○ Asexual reproduction occurs, but sexual reproduction is more common
○ Most species are gonochoristic
○ No permanent sex organs
■ When the worm is mature, it is packed with sperm and eggs
○ External fertilisation
○ Eg
...
sea mouse
● Chaetae scatter light, creating patterns of colours
■ Live beneath stones, in coral crevices, and in algae
○ Pelagic dwellers
Joanna Griffith (2017)
○
○
■ Similar to surface dwellers, but transparent
Burrowers
■ Occupy vertical or U-shaped burrows
■ Prostomial sensory appendages are generally absent
Tubicolous
■ Live in tubes, tubes may be composed of secreted material
Oligochaetes
● Eg
...
crown of thorns), oyster beds
(eg
...
orange sea cucumber
● respiration/circulatory system/excretion
○ Unique respiratory tree
■ Branching tube inside anus for gas exchange
● Trunks of trees merge with the cloaca
● Water circulates through the tubules by the pumping action of
the cloaca and respiratory tree
● Gases are exchanged with the coelomic fluid
● Defence
○ Cuvierian tubules discharge
■ Long, sticky, toxic threads
■ Tubes of Cuvier regenerate
○ The most extreme defence mechanism is evisceration
■ Discharge of digestive tract, respiratory tract, and gonads
■ May also be used to purge the gut of waste
Crinoidea
● Primitive sessile class (similar to 500 million year old fossils)
● Sea lilies, feather stars
● Pentaradial symmetry
○ Five or more flexible arms
● U-shaped gut, anus next to mouth
● No madreporite, pedicellariae, and spines
● Suspension feeders
Concentricycloidea
● Primitive sessile class
● Sea daisies
● First discovered in 1986 in New Zealand
○ Three species in the Bahamas and North Pacific
○ Relationships to other Echinoderms is unclear
● No arms
● Pentaradial symmetry
● Tube feet on the periphery of the disc
● No anus, no intestine
--------------------------------------------------------------------------------------------------------------------------Joanna Griffith (2017)
13,15,16,17,18,&19: ARTHROPODA
Most invertebrate species are arthropods
Ecdysozoans
○ Animals that shed an external ‘coat’ (cuticle) when they moult (ecdysis)
○ Most successful and abundant animal group
Body plan
● All arthropods have a fairly similar body plan
● Exoskeleton
○ Chitinous exoskeleton or cuticle
■ Cuticle is soft until it is sclerotised
■ Cuticle is divided into several plates
● Plates are connected by articular membranes
○ Many arthropods also have articular condyles and
sockets (eg
...
in arachnids)
● Malpighian tubules
○ Involved in the removal of nitrogenous waste from the blood in terrestrial taxa
○ Lie in hemolymph alongside the gut in arachnids and hexapods
○ Secrete uric acid into the hindgut
● Aquatic taxa use saccate nephridia that secrete ammonia, typically across the gills
○ Ammonia is toxic and must be diluted in water during excretion
■ Terrestrial taxa secrete uric acid, which is less toxic and therefore
does not require as much water
Nervous system/sense organs
● Like other animals, the basic units of a neuron are the dendrite and axon
○ Neurons are interconnected to form nerve centres (ganglia)
● Brain
○ Anterior protocerebrum
○ Medium deutocerebrum
○ Posterior tritocerebrum
● Arthropod nervous systems are a major target for insecticides
○ DDT and pyrethroid: target sodium channels in axons, blocked channels
cause paralysis
○ Organophosphates: target acetylcholinesterase, the deactivated enzyme
means that neurotransmitters are not removed from the synaptic cleft
○ Neonicotinoids: target acetylcholine receptors, blocking the receptors means
that the neurotransmitter cannot function
● Sense organs
○ Having an exoskeleton can make it more difficult to sense the environment
○ Sensilla
■ Have chemo- or mechano-receptors that detect external stimuli
○ Eyes
■ Vary in complexity
● Can be simple eyes with a few photoreceptors or complex
eyes with thousands of photoreceptors
■ Images produced are crude
■ Poor distance vision
■ High flicker-fusion rate
■ Eg
...
pheromones in moths (fluffy antennae)
Locomotion
● Musculature
○ Striated muscle bundles
○ Not attached directly to the hard part of the exoskeleton, instead attached to
the epidermis, which is then attached to the hard cuticle by microtubules
■ Allows the organism to still use its muscles to escape predators after
moulting
○ Specialised for rapid movement
● Movement
○ The muscles and skeleton work together
○ Typically, muscles work in pairs (extensors and flexors)
○ Increase in blood pressure can extend plates in the thorax/abdomen
○ Generally, jointed appendages are used
○ Speed of movement is dependent on appendage length
○ Gait
■ Wave of leg movement
● Legs on opposite sides of the body alternate
● Undulation
○ Efficient locomotor-skeletal system
■ Tubular structure resists bending
■ But, if the animal were to grow too large, the soft exoskeleton would
buckle after moulting
Key adaptations for life on land
● Hard exoskeleton with waxy epicuticle
○ Small organisms have a large surface area to volume, so the exoskeleton
helps to trap water
● Jointed limbs with opposing muscles
● Efficient nitrogenous waste excretion via uric acid
● Gas exchange systems that can function in air (trachea or book lungs)
● Hindgut to recover water from food
● Internal fertilisation
● Pre-adaptation: a shift in the function of a trait during evolution
○ A trait may evolve because it serves one particular function, but subsequently
it may come to serve another
Trilobitomorpha
Joanna Griffith (2017)
Antennae, legs
10 orders, 150+ families, 5,000 genera, 20,000 described species
○ Useful for dating rocks
● Trilobite means ‘three-lobed’
○ Describes body segments
● Modern arthropods
○ The earliest fossils with an arthropod-type body plan are from the Cambrian
explosion
■ Body segments were equal to each other
● Through evolution, the segments became specialised
Chelicerata
● Body plan
○ Clawlike appendages called chelicerae
○ Cephalothorax and abdomen
○ No antennae
○ Simple eyes (single lens)
○ Chelicerae, pedipalps, legs
● Eurypterids
○ Sea scorpions
○ Extinct in Permian period
■ Ancestors of arachnida?
○ Top marine predators (1-2ft long)
● Xiphosura
○ Most are extinct, only horseshoe crabs remain
■ ‘Living fossils’
■ Characteristic trisegmented chelicerae on either side of the upper lip
■ 5 pairs of walking legs, posterior to the chelicerae
● Bases of legs have gnathobases, which aid in food processing
■ Blue blood
● Contains copper (hemocyanin) rather than iron
● Arachnida
○ Includes scorpions, spiders, harvestmen, mites, and ticks
○ Body plan
■ Probably evolved from aquatic environments
● Epicuticle became waxy to reduce water loss
● Gills became modified for use in air (book lungs and trachea)
● Appendages became better adapted for terrestrial locomotion
■ Poison and silk glands
○ nutrition/digestion
■ Most arachnids are carnivorous
■ Prey are captured and killed using the chelicerae
■ Food is rapidly digested externally
● Food ‘broth’ is sucked up into the prebuccal cavity
○ respiration/circulatory system/excretion
■ Book lungs (invaginations)
● The book lungs in Xiphosura are exvaginations
●
●
Joanna Griffith (2017)
Trachaea and malpighian tubules evolved independently twice in
arthropods
Order Scorpiones
■ Among the oldest terrestrial arthropods
■ Generally nocturnal
● Fluorescent under UV light
■ Relatively large, can be between 3-9cm
■ Body plan
● Body consists of a cephalothorax covered by a carapace, a
long abdomen, and a sting
○ In the middle of the carapace are a pair of large median
eyes
● Small anterior chelicerae and enlarged pedipalps that form
pincers
● The abdomen has 7 pre-abdomen segments and 5
post-abdomen narrow segments
● An opercular plate covers the genital opening
● Posterior to the opercular plate are sensory pectine near the
ventral midline
■ nutrition/digestion
● Most species are sit and wait predators
● Use their stings and/or pedipalps (pincers)
■ reproduction/development
● Males initiate courtship
○ Courtship may take minutes or hours
● Male deposits a spermatophore on the ground and positions
the female over it
● All scorpions brood eggs in the female reproductive tract
○ Aplacental viviparous species
● Development takes place in the ovaries
● After up to a year, 1-90 offspring are produced
Order Thelyphonida
■ Whip scorpions
■ Eg
...
compound eyes, colour vision (jumping
spiders)
■ Silk
● Each spinneret has many openings from the silk gland
● Silk is a protein, secreted as a liquid
● Many uses for silk, eg
...
25-0
...
Eriophyidae
○ Family of 200+ genera of mites
○ Worm-like, only have 2 pairs of legs
○ Chelicerae are modified into needle-like stylets to
pierce and suck out plant cell contents
○ Eg
...
Varroa destructor
○ Parasitic mite that attacks honey bees and causes the
disease Varroatosis
■ Female mites enter the honey bee brood cells
and lay eggs on the larvae
● When the larvae hatch, they suck the
bee larvae hemolymph, weakening them
● Eg
...
Ixodes
○ Lyme disease, carried by a bacterium belonging to the
genus Borrelia
■ Carried by ticks in the Ixodes genus
● Tick has a 2-year lifestyle
● As it feeds, it transmits the disease
● Eg
...
Illacme plenipes has 750 legs
● respiration/circulatory system/excretion
○ Gas exchange is through a tracheal system
○ Spiracles are always open, leads to water loss
○ The heart is a dorsal tube that extends through the body
○ Have hemocyanin (copper instead of iron)
● Nervous system/sense organs
○ Nervous system isn’t centralised
● Chilopoda
○ Centipedes
○ Most North American and European species are 3-6cm long, tropical species
can be up to 30cm long (Scolopendra gigantea)
Joanna Griffith (2017)
Temperate zone centipedes tend to be reddish brown, but tropical species
can be any combination of colours
○ Body plan
■ Antennae on the front margins
■ Mandibles bear teeth and a fringe of setae
■ Covering the mandibles are a pair of forcipules (poison claws)
● At the terminal end of each forcipule is a fang
○ nutrition/digestion
■ Predators, many eat small arthropods
■ Prey is detected with the antennae
■ Prey is killed or stunned with the forcipules
● The neurotoxic venom is often painful but not always deadly to
humans
○ reproduction/development
■ Females have one ovary, and males have one or more testis
■ Females and males have a gonopod which aids in handling of the
spermatophore
■ Sperm transfer is indirect (via a spermatophore)
■ Some centipedes brood their eggs in clusters of 15+
■ The female winds her body around the eggs or carries the eggs
between her gonopods for a short time before depositing them
○ Locomotion
■ Centipedes that are adapted to run have longer legs
■ Centipedes that have adapted for burrowing through soil and humus
have short legs that anchor the body
■ Eg
...
by tapping, drumming,
releasing pheromones
■ The eggs are fertilised from stored sperm and laid (up to 300 eggs at a
time)
● Some deposit eggs in soil or humus, others construct a “nest”
○ Locomotion
■ Most move slowly
■ Adapted for pushing through humus and leaves
■ The backward, pushing stroke is activated in waves along the length
of the body
■ pushing/burrowing is facilitated by the body plan
○ Defence
■ Compensate for slow gait with a calcareous exoskeleton
■ Some can roll up into a ball
■ Use repugnatorial glands to repel small animals
■ Bioluminescence may result in some millipedes being attacked less
Symphyla
○ Small class (about 160 species)
○ Live in soil and leaf mould, in most parts of the world
○ Very small, only about 2-10mm long
○ Body plan
■ Trunk has 12 leg-bearing segments
■ The last segment carries a pair of spinnerets or cerci (usually act as
sensory organs) and long sensory hairs
■ Mouthparts are similar to insects
○ reproduction/development
■ Genital opening is on the ventral side of the 4th segment
■ Male deposits 150-450 spermatophores on stalks, and the female
finds and eats them
○ Locomotion
■ Very fast, can twist and loop their bodies as they move through the
humus
● Probably an adaptation to avoid predators
Pauropoda
○ Fairly abundant class (about 380 species)
○ Widespread in temperate and tropical regions
Joanna Griffith (2017)
○
○
○
○
Minute, about 0
...
in the southern ocean, dust blowing in from islands feeds plankton
blooms, increasing crustacean populations, increasing prey for many marine
mammals and birds
● Body plan
○ Two pairs of antennae (defining feature), mandibles, two pairs of maxillae
○ Highly specialised appendages
○ Walking legs on thorax and abdomen
○ Trunk anatomy
■ Composed of distinct and similar segments, and a telson
● Basic design is often modified by the fusion of segments
■ In some groups, the trunk is divided into the thorax and abdomen with
various numbers of segments
■ The thorax or anterior trunk segments are covered by a dorsal
carapace
● Often, the lateral margins of the carapace overhang the sides
of the body
○ Appendages
■ Biramous (divide to form two branches)
● There is a basal protopodite composed of a coxa and basis
○ Cuticle
■ Calcified (in contrast to most arthropods)
■ Epicuticle and procuticle contain deposits of calcium salts
■ The outer layer of the procuticle is pigmented and contains tanned
proteins
● nutrition/digestion
○ Large range of diets and feeding mechanisms
■ Eg
...
woodlice)
■ Have gills, which must remain moist
Hexapoda
● Includes animals that:
○ Shed an external cuticle when they moult (ecdysis)
○ Have an exoskeleton
○ Have a three-part segmented body
○ Have six (hex) jointed, paired appendages
● Body plan
○ Amalgamation of segments into functional units (tagmosis)
■ Three main regions called tagmata
○ Huge diversity of morphology from the same basic body structure
● Insecta
○ Largest group (about 750,000 species)
○ Very successful
■ Occupy most environments
■ Very ecologically important
○ Body plan
■ One pair of antennae, one pair of mandibles, two pairs of maxillae,
three pairs of legs
■ Three body sections (tagmata)
● Thorax is split into pro-, mesa-, as well as notum/tergum,
sternum, and pleurons
Joanna Griffith (2017)
○
○
○
○
● Abdomen is made up of 9-11 segments
■ Usually, there are two pairs of wings on the thorax
■ One pair of compound eyes
nutrition/digestion
■ Adapted to many diets
■ Modifications of mouthparts reflect how food is ingested
● Mandibles function in cutting and tearing
● Maxillae and labium function in food handling
● Piercing mouthparts
● Sucking mouthparts (eg
...
horseflies)
● Sponging mouthparts (eg
...
Cellembola (springtails), Zygentoma/Thysanura (silverfish)
Joanna Griffith (2017)
exite/endite hypothesis
● Wings evolved from a gill branch and a leg segment
● Eg
...
dragonflies)
● The evolution of sclerites has enabled insects to fold the wings
over the abdomen
■ Insect flight muscles are very powerful
● Huge mitochondria within the muscle cells
● At low body temperature, metabolic rate is low, so many flying
insects have to ‘warm up’ to enable flight
Coevolution of plants and insects
■ Two interacting partners change in relation to each other
■ Eg
...
strong selection placed on the ability to produce cyanide
for defence
■ Implies reciprocal linked interactions, which are difficult to prove
■ The evolution of flowering plants created a huge diversity of new
niches for insects
● About 50% of insects are herbivores, but only 9/30 orders are
primarily plant-eaters
Order Coleoptera
■ Beetles
■ “Sheath wing”
■ Contains more species than any other insect order
● 40% of all insects are coleoptera
● 25% of all species are coleoptera
● 400,000 named species, but estimates go up to 8 million
■ Older than the dinosaurs (the first “true” coleoptera appeared 230mya,
in the Triassic period)
■ Body plan
● Coxae (first leg segments) are housed in a special cavity,
which makes beetle bodies very compact
○ Makes hind coxae immovable
● Elytra
○ Modified forewing
○ Protects the wings and body from damage and infection
(eg
...
6mph
Eg
...
mealworms are capable of degrading polystyrene
into usable organic matter at a rate of about 35-40mg
per day
■ reproduction/development
● Some species (eg
...
leaf beetles (family Chrysomelidae)
○ Over 37,000 species in over 2,500 genera
● Beetles were among the first insects to visit flowers
○ Remain essential pollinators, especially for ancient
species such as magnolias and spice bush
Order Hymenoptera
■ “Married wing”
■ Known for extreme cooperation
● Build huge nests
■ Body plan
● Large diversity of sucking/chewing mouthparts
● Hind wings are connected to forewings by a series of hooks
● “Nipped waist” due to fusion of first abdominal segment to
thorax
■ Social complexity
● Hymenoptera contains the largest group of eusocial insects
○ Some are only primitively social, eg
...
queens, soldiers, workers
● Basic modifications to the body plan promote task
specialisation
○ Tending eggs and young
○ Providing for the colony (trophallaxis, the social sharing
of liquid food - also used to pass chemical signals
among each other and maintain a cohesive colony
identifying odour)
○ Defending the colony
■ Soldiers may have massively distorted
defensive structures, which may result in them
losing the ability to feed themselves
■ Eg
...
wax moth larvae (Galleria mellonella)
○ Eg
...
vampire moth (Calyptra minuticormis)
● Can be highly damaging pests
○ Four lepidoptera species are in the top ten
insecticide-resistant insect species
■ reproduction/development
● Holometabolous (complete metamorphosis)
● Mating and egg-laying occurs on a host plant
● Larvae are grub-like
■ Colouration
● Eyespots can deter vertebrate predators (proven to drastically
reduce mortality)
● Many species feed on toxic plants (eg
...
Heliconius melpomene and
Heliconius erato
■ Batesian mimicry: non-toxic species copying a
toxic species
● Only works if the toxic species is
common and the mimic species is rare
Order Diptera
■ flies
■ Highly variable ecology
■ Body plan
● Functional forewings
○ Rapid and agile flight in some species
○
○
Joanna Griffith (2017)
Variable mouthparts and body form
Halteres
○ Reduced hindwings
○ Beat in time with forewings
○ As the body turns in flight, gyroscopic forces are
exerted on the halteres
■ Sensitive hairs sense these forces
■ Allow the fly to make adjustments and stabilise
the body rapidly during flight
nutrition/digestion
● Some herbivorous species
● Many parasitic species
● Most feed on decaying organic matter
reproduction/development
● Holometabolous development (complete metamorphosis)
pests/disease vectors
● Order contains many of the most important disease vectors
○ Eg
...
Culex pipiens
■ Wings clear/membranous
■ Short palps
■ Body horizontal when feeding
■ Transmits encephalitis and filariasis
○ Eg
...
Cochliomyia hominivorax (the new-world screwworm)
○ Larvae feed in open wounds and living flesh
○ Infested wounds attract more oviposition
■ Thousands of eggs in one wound can kill cattle
within 7-10 days
○ Alternative hosts include humans and wild animals
○ Sterile insect technique (mass sterilisation and release
of male individuals) has been successful in reducing
the population
■ Barrier to reinvasion maintained by releasing 50
million flies per week
● Flies can also be beneficial to humans
○ Eg
...
Tachinidae
■ Lay eggs on top of common pest insects, larvae
feed off the host and eventually kill them
● Biological control of winter moths in
North America
○ Pollinators
○ Waste disposal
■ Eg
...
Scathophaga stercoraria (yellow dung fly)
■ Eg
...
Strongyloides (threadworms)
Eg
...
Dictyocaulus viviparus (lungworms)
○ Vertebrate parasites
● Eg
...
Myrmeconema neotropicum
○ Induces fruit mimicry in the tropical ant Cephalotes atratus
Body plan
● Cylindrical bodies
● Body is covered by a tough cuticle, which they moult
● No circulatory system
○ Nutrients are transported via fluid in the pseudocoelom
● Body wall muscles are longitudinal
○ Contractions produce a thrashing motion
● Can infect humans and animals
Joanna Griffith (2017)
Title: 1st: Introduction to Invertebrate Zoology
Description: 1st year Introduction to Invertebrate Zoology notes, University of Exeter
Description: 1st year Introduction to Invertebrate Zoology notes, University of Exeter