BIOL 321 Part II Flashcards
Bryozoa lifestyle
colonial, sessile, suspension feeders
Individual bryozoan
zooid
very tiny (mm’s)
lophophore
lophophore
funnel of ciliated tentacles used for suspension feeding
Bryozoan exoskeleton
zooecium
often calcified
Bryozoan digestion
u-shaped digestive tract
why is u-shaped digestive tract appropriate for exoskeleton enclosed organisms
if posterior end is closed feces would accumulate
Bryozoan compartments
2 coelomic compartments - lophophoral coelom, perivivsceral coelom
Bryozoan organs
no specialized organs for gas exchange, excretion/osmoregulation, internal fluid circulation
How do Bryozoans accomplish gas exchange/fluid circulation without organs
ciliated mesothelium circulate fluids/gases
Bryozoan colony form diversity
stoloniferous
encrusting
erect
encrusting bryozoan colonies
zooids lie flat on substrate w/ dorsal surface attached to substrate
stoloniferous bryozoan colony
creep over rocks, zooids bud up from stolon
erect Bryozoans
flat blades
branched fronds
solitary bryzoan
Monobryozoan
solitary, motile, infaunal- interstitial fluids of sand grains, extensions hold on to sand grains, contact to pull organism down
Bryozoan polymorphs
heterozooid
autozooid
Heterozooid
usually defensive, non-feeding
e.g. avicularium, operculum snaps shut to deter predators/organisms settling
Autozooid
feeding, reproducing
Vibraculum
type of heterozooid (Bryozoan)
elongate ‘whips’ swarm around colony, dislodge settling/predators/used as ‘legs’ for movement
Bryozoan reproduction
larva -settles -metamorphosis -initial zooid (founder)- aseuxal budding - more individuals = colony
sexual reproduction to form new colony
Bryozoan circular colony
founder cell in middle
budding takes place around circumference
periphery of encrusting bryozoan colony
budding zone
brown body
degenerated mass of zooid tissue, only epidermis +mesothelium, regressed zooid
what happens with brown body
regenerate whole new zooid w/ brown body in stomach - defacate it out
why do zooids regress and regenerate?
possibly method of dealing with toxin/waste build up
Bryozoan defense
heterozooids colonialism chemical defense induced defences calcification of frontal membrane
colonial defense
strength in numbers, genome maintained - unlikely predator will eat whole colony, can regenerate
what are chemical defenses
secondary metabolites
deter predators
antibiotics
Bryozoan secondary metabolites
bryostatin - appears to have anti-cancer, anti-alzheimer properties, synthesized by symbiotic bacteria
why is it important to calcify frontal membrane of Bryozoan
flexible frontal membrane leaves them vulnerable to predation
e.g. nudibranch slices frontal membrane w/ radulae and sucks out bryozoan
Induced defense, bryozoa
initially put all E into reproduction, predator comes, w/i 48hrs of predator feeding zooids form calcareous spines on corners of zooecia
Bryozoan frontal membrane calcification
frontal surface inflexible to protect against predators
how can lophophore retreat with inflexible frontal membrane
sac inside of perivisceral compartment is filled w/ fluid, ejects fluid out pore to make room for lophophore
periversceral fluid-containing sac in Bryozoans with frontal membrane calcification
Ascus
how is lophophore moved out in Bryozoan w/ frontal membrane calcification
parietal muscles expand balloon and water is sucked back in - forces lophophore out of way
how is lophophore retracted in a Bryozoan
lophophore retractor muscles
Bryozoan sexual characteristics
gametes arise from mesothelium
hermaphroditic (sequential or simultaneous)
sperm spawn through tentacular pores
external or internal fertilization
Bryozoan external fertilization
both gametes spawned externally
long-lived feeding larvae
Bryozoan internal fertilization
eggs maintained internally, brooded for short period, short-lived non-feeding larvae
Bryozoan egg brooder
ovicell (heterozooid)
Mollusca subphyla
Conchifera
Aculifera
Aculifera groups
Polyplacophora
also Chaetodermomorpha, Neomeniomorpha which we are not studying and are placaphora
Conchifera groups
Monoplacophora Cephalopoda s.g. Scaphopoda s.g. Bivalvia s.g. Gastropoda
Mollusca primitive characteristics
dorso-ventral differentiation
CaCO3 shell
mantle cavity, gills, osphradia
shell-attached muscles
chiton shell
8 articulating shell valves w/ 8-pairs dorsal ventral shell-attached muscles (from shell-foot)
Mollusc dorso-ventral differentiation
visceropallium
cephalopodium
Chiton osphradia
2 sensory organs - on either side of anus
Mollusca gill
ctenidium - central axis w/ gill lamella, afferent/efferent blood vessels
Mollusca circulation
hemal fluid in to afferent blood vessel, through each gill lameli, out efferent blood vessel
Mollusca water circulation
in between each lameli - countercurrent to hemal flow to maximize efficiency of gas exchange
Mollusca reduced coelom
pericardium
containing heart, attached to metanephridia, gonads, associated with gut
metanephridia, Mollusca
heart contraction = ultrafiltration
fluid down metanephridial ducts = selective reabsorption
Mollusca gonads
derived from mesothelium
most extant organisms do not retain connection between gonad-pericardium
Mollusca digestion
mouth, radular cartilage, foregut, radular sac
radular sac
secretes riot of radular chitinous teeth on ribbon
radular cartilage
rods, support radular teeth
how radula works
muscle protrudes radular cartilages out of mouth carrying radular teeth - pull teeth back in scraping substrate
Mollusc radula + cartilages
buccal mass
Mollusc nervous system
circum-esophogeal nerve ring (cephalopodium) visceropallial nerve cords pedal nerve cords pleurovisercal ganglia cerebral ganglia pedal ganglia
distinctive characteristics of chitons
minimal cephalization dorso-ventral flattening dorsal shell w/ 8 articulating valves mantle assisted substrate adhesion radula with magnetite caps
Chiton cephalization
non-ganglionated
non-active, head non-specialized
Why chitons have dorsoventral flattening
low profile helps avoid being washed away - adaptation to wave swept shores
Chiton substrate adhesion
very muscular foot + mantle periphery for clinging
lift mantle roof = negative pressure = suction
Chiton radula caps
Mollusc teeth are replaced but chitons cap w/ magnetite (Fe containing bxomineral) to reduce wear
Chiton reproduction
dioecious, broadcast spawn (m and f), external fertilization, gametes from gonad mesothelium, ciliated non-feeding larvae
largest metazoan phyla
Arthropoda
2nd largest metazoan phyla
Mollusca
Class Monoplacophora
very rare, deep sea, only known from fossils, single dorsal shell, ventrally similar to chiton, sister clade to rest of conchifera
Monoplacophora ventral
mouth, anus, lateral mantle cavity w/ ctenidia, 8 dorsal shell muscles
Gastropod shell
isometric coiling -coils in same plane, only see one from front view -asymmetric coiling
Accommodating coiled shell, gastropod
visceroplallial elongation and coiling
gastropod shell central pillar
columella, CaCO3, central coiling axis
shells are designed to be
protective retreat
gastropod shell-attached muscles
single or single paired
hold on to columella
run down to foot/head to pull head in, close operculum
changes from monoplacophoran to hypothetical intermediate gastropod
Shell coils (exogastric) and elongate dorsoventrally, reduced shell aperture, mantle cavity, # shell muscles, #ctenidia
changes from hypothetical intermediate gastropod to derived gastropod
Torsion (180ºrotation) of visceropallium relative to cephalopodium, rotation of shell (endogastric shell coil)- anus + mantle cavity over mouth
evidence of torsion
anatomy of living gastropod
development of basal gastropod
asymmetries in gastropods
- asymmetric coil of shell (goes to the left or right)
- torsion
- non-bilateral organs/structures (majority of gastropods have left ctenidium/osphradium)
extant gastropod anatomy as evidence of torsion
cross-over of visceropallial nerve connectives
gastropod veliger larva
tiny calcareous shell swimming side down, 2 velar lobes, modified trochophore (prototroch, metatroch for feeding)
torsion advantages
mantle cavity and therefore ctenidia are moved anterior - water that is brought in is less disturbed by movement
torsion advantage to developing gastropod
mc below foot = pull foot in first, velum in last; velum is more valuable and vulnerable than foot, advantageous to have mc above head and pull foot in last
development of basal gastropod as evidence of torsion
ontogenetic torsion
see foot on both sides of shell in developmental stage
torsion hypothesis test, abelone
1 batch pre-torsional larvae, 1 post-torsional; both have predators; found no difference in # survivors- no evidence of torsion being advantageous
1 problem of torsion
results in deeper anterior mantle cavity - water needs to be frequently circulated for aeration and removing waste
solutions to fouling
restricted mantle cavity
shell perforations
shit of anus to right side
all gastropods with shell perforations
have 2 ctenidia
shifted anus, gastropod
Majority of gastropods; loss of right ctenidium, osphradium;
shift anus to right;
oblique current through mantle cavity; pick up fecal material last on way out
gastropod snorkle
siphon
right side, highly mobile, muscular, samples water and picks up chemical signatures
alternative hypothesis for gastropod body
unilateral enlargement of mantle cavity- 2 mc’s and one expands in different ways
types of gastropod MC enlargement methods
monotocardian
diotocardia
heterobranchia
monotocardian
enlarged MC, ctenidia on left, anus on right
diotocardian
enlarged MC, 2 ctenidia (L/R), anus in middle
heterobranchia
MC not enlarged, reduced to 1 ctenidia
gastropod cephalization
fast moving, A-P axis = need for receptors, ganglionization: 1+ tentacle sets, eye spots, series of ganglia
Class Gastropoda main features
protective shell - single plate into coiled cone
torsion
pronounced development of head
Class Gastropoda groups
Patellogastropoda s.g.
Vetigastropoda s.g.
Caenogastropoda s.g.
Heterobranchia
Old Class Gastropoda phylogeny
Prosobranchia (Patellogastropoda, Vetigastropoda, Caenogastropoda) s.g. Opisthobranchia (Heterobranchia) s.g. Pulmonata (Heterobranchia)
Vetigastropoda
keyhole limpets, abelone
Caenogastropoda
majority of marine gastropods, well developed shells
Gastropoda primitive feeding method
herbivorous grazing using radula
radula
ribbon of teeth secreted by radular sac, protruded out of mouth to scrape rock
buccal cavity
area of gut that radula opens in to
vetigastropod feeding
many herbivorous grazers, retain primitive method
Caenogastropod feeding
many herbivorous grazers, retained primitive feeding method (radula); many predators with proboscis
Proboscis
right side, normally tucked in, used for feeding
herbivorous gastropod foregut
mouth - buccal cavity- anterior esophagus- mid-esophageal gland; salivary glands attach to buccal cavity
predatory gastropod foregut
deep in-pocket in anterior end = proboscis sac, buccal cavity = proboscis, esophagus greatly lengthened, mid-esophageal gland is enlarged and connected to esophagus by narrow gut
evidence of predatory gastropods
shell drills, leave bevelled edged hole on preys shell
how predatory gastropod makes shell drills
tip of proboscis has accessory boring organ - raps on shell = mechanical abrasion; also chelate shell = chemical dissolution
Cone snail
highly derived gastropod feeding, mostly tropical/subtropical, predatory caenogastropod, feeds on worms/molluscs/fish
Cone snail feeding
very long proboscis, w/ harpoon tooth and venom
cone snail harpoon tooth
apex of radular tooth is shaped like hollow harpoon, snail takes 1 tooth and places it at end of proboscis, tooth is connected to venom gland
cone snail venom gland
midesophogeal gland highly elongated ending in muscular ball specialized to synthesize and secrete conotoxins
conotoxin
neurotoxins, peptide that bind to Fe channels and neurotransmitters, rapidly immobilize prey, any 1 species may have 100’s
Heterobranchs
elaborated dorsal surface, secondarily deflected anus to posterior end (detorsion); loss of larval shell during metamorphosis
Heterobranch loss of shell
do have shell in juvenile form (and torsion), crawl out of shell and discard it and operculum
Heterobranch defense
chemical defense
escape behaviour
sequester nematocysts
autotomy
chemical defense, Heterobranch
dorsal chemical glands - unpallatable/toxic
where do Heterobranchs get chemical defense
some make own
many steal from prey (sponges, bryozoans) and put in dorsal gland sacs (cerata/papillae)
escape behaviour, heterobranch
touched by predator – lift off seafloor - d/v body undulations - catch current and float away
sequestering nematocysts, Heterobranchia
feed on cnidarian- sequester nematocyst- carried up to cerata and deposited- phagocitized - utilized
where are nematocysts stored in heterobranch
cnidosac - special sac at time of cerata lined by epithelium
how are heterobranchs able to utilize nematocysts?
secrete mucus that makes them not trigger (like clown fish) - may get stung at first until they learn the right formula (species specific)
Nudibranch autotomy
rapidly release body part, mainly cerata
terrestrial heterobranchs
snails, slugs; terrestrial; threats include desiccation, T changes
terrestrial heterobranch, breathing
closed mantle cavity = ‘lung’, pneumostome = pore, opening to lung (able to close)
adaptations for life on land (Heterobranchia
mantle cavity forms internalized lung
conversion of ammonia to uric acid
asetivation
Aestivation, heterobranchs
metabolic slow-down, hide in humid location,
tolerance of desiccation
ability to rapidly rehydrate
freshwater snail w/ air breathing adaptation
Caenogastropods, pneumostome + long siphon opening in to lung
(convergence w/ heterobranchs)
how gastropods can rehydrate
open spaces between epithelial cells
Vetigastropoda reproduction
broadcast spawn gametes, external fertilization
Caenogastropoda, Heterobranchia reproduction
internal fertilization, encapsulated eggs, juvenile crawls out of egg capsule
Class Bivalvia
dv flattened body 2 shell valves spacious lateral mantle cavity to house ctenidia minimal cephalization no radula
Bivalve falttening
including foot, facilitate digging
why bivalves need 2 valves
life in substrate - need to remain open for circulation, need to shells to remain open - pressure would collapse mantle
Bivalve radula
none in extant members, feeding by suspension or deposit
bivalve water flow
along edge of shell or restricted to siphons
bivalve adductor muscles
pull valves together, cause tension in ligaments, when released ligaments pull shells open
Bivalve shell secretion
secreted by outer mantle lobe, sequential layers of bxomineral CaCO3 w/ crystals oriented in opposite directions - cross-hatched for strength
Bivalve mantle margin
sensory structures on periphery of mantle fold (middle lobe)
bivalve middle lobe
most exposed to environment, may contain photoreceptors, tentacles
bivalve outer lobe
secretes shell
bivalve inner lobe of mantle fold
‘muscle lobe’
has pallial muscles connected to shell valve to pull and tuck all soft tissues in side shells
Diversity of feeding in Bivalvia
prosobranch
lamellibranch
septibranch
Protobranchs
deposit feeder; use palp tentacles; gills for gas exchange only, ancestral feeding strategy, long paired siphons
palp tentacles
ciliated, deposit feeding in protobranchs
Protobranch ctenidium
gas exchange only
non-elaborate gill filaments on sides of gill axis
Lamellibranch
ctenidia for gas exchange and feeding, suspension feeders
lamellibranch ctenidia
elongate filaments, folded back to fit, highly ciliated to carry particles
lamellibranch folds
demibranchs
outer demibranch = closest to shell
Lamellibranch cilia
each ‘limb’ is surrounded by 3 types of cilia: lateral cilia, laterofrontal cilia, frontal cilia
lateral cilia, lamellibranch
short, on the ‘top/bottom’, create water current
laterofrontal cilia, lamellibranch
longer, point diagonally, intercept particles
Septibranch feeding
rare, predators, suction feeding, enormous inhalant siphon used for sucking in small organisms
septibranch ctenidia
modified gill = muscular, perforated diaphragm; closed shell, elevated muscular shelf = increased volume in inhalent chamber = suction
Bivalvia habitatlifestyle diversity
burrowers (majority; deeper = longer siphon)
attached to solid substrate
boring
example of shallow burrowing bivalve
cockle
e.g. of bivalve attached to solid substrate
mussels
Mussel attachment
secrete byssal threads
frontal cilia, lamellibranch
short, face out, frontal surface, carry particles down filament to elbow-like area of gil filament
example of deep burrowing bivalve
geoduck
how to use byssal threads
byssal gland by foot secretes fluid– fluid runs down foot, forms puddle– starts to harden - lift foot- move foot and repeat
oyster substrate attachment
cement w/ CaCO3
boring bivalve
shipworms
free-swimming larva, settle, metamorphose, feed/create tunnel
Class Scaphopoda
tusk shells, see text book ch 12
Class Cephalopoda
active, pelagic, predatory, smartest, largest, fastest molluscs, 3 subclasses, 1 extinct
Cephalopod molluscan characteristics
radula
molluscan-style gill (but not ciliated)
shell-secreting mantle (but reduced/lost in most extant)
Cephalopod unique characteristics
septate shell
highly modified foot
Cephalopod foot modifications
prehensile appendage (arms, tentacles) funnel
Cephalopoda groups
Nautiloidea - extant, since Palaeozoic, first known group (540Ma)
Ammonoidea - extinct end of Mesozoic, known from mid Paleo
Coleoidea - extant, known from end of Paleo
Nautilus
primitive cephalopod, tentacles have no suckers, external shell, hood, funnel
Nautilus shell
external, gas-filled chambers, chambers separated by septa, septa perforated for siphuncle
Squid
active, agile swimming, predatory feeding
squid example
Loligo
squid predation
visual predators- image-forming eyes, eye morphology convergent w/ vertebrates
Coleoidea metabolics
high metabolic rate, ventilate gills by pumping muscular mantle, closed circulatory system, systemic and branchial hearts
Coleoid branchial hearts
booster pumps to force blood through gills
Protostomia groups
Lophotrochozoa
Exdysozoa
Exdysozoan groups
Nematoda, Onychophoran, Arthropoda
Ecydsozoa shared morphological characteristics
Ecdysis
no motile cilia/flagella
Ecdysis, ecdysozoa
periodically moult exoskeleton/cuticle for growth
Phylum Nematoda basic characters
multilayered collagenous cuticle, 4 moults, longitudinal muscles only, pseudocoel, syncytial epidermis, anterior nerve ring, longitudinal cords, aberrant cilia, eutely
Nematode cilia
non motile, restricted to sensory function
eutely
embryo hatches w/ set # of somatic cells and never produces any more; growth only by enlargement of cells
Nematode body movement
thrashing motion; dorsoventral contractions; only useful for forward motion in dense medium
Needed for nematode locomotion
stiff cuticle;
fluid maintained under high pressure in pseudocoel;
longitudinal muscles
Nematode feeding
triradiated pharynx, muscle contraction opens lumen, have to actively open gut by muscular means b/c high pressure of gut, must drink water to actively push food down
Nematode reproduction
Asexual and sexual
nematode asexual reproduction
very rare - parthenogenesis (no budding or fission)
nematode sexual reproduction
dioecious, internal fertilization, males w/ copulatory spicules, ameboid sperm (aflagellate)
nematode developments
4 moults to adult stage; egg - 4 juveniles - adult; eutely
Dauer larva
Nematode facultative diapause, triggered by enviro. cues, age arrest (reduced metabolic rate), occurs at L2 (juvenile #2)
parasitic nematode examples
hookworm, wuchereria, golden nematode
Hookworm
intestinal parasite, 1 host, cuticle around mouth forms tooth/hook-like projections, consume blood from intestinal wounds, leads to anemia
Hookworm life cycle
adult worm in human intestine - eggs passed in feces - juv. 1 hatches - 2 moults - juv. 3 burrows into skin, often foot - moves to circulatory system - heart - lungs - trachea - pharynx– intestine
Wuchereria bancrofti
Nematode, Elephatiasis, microfilariae larvae clog lymphatic vessels, causes grotesque swelling, 2 hosts - human, mosquito
Wuchereria bancrofti life cycle
microfilariae - mosquito - moult - move to salivary gland - transmitted to human
Golden nematode
1 host, damaging potato parasite, cysts on roots are dead swollen fm bodies filled w/ eggs
Golden nematode distribution
Europe, Asia, Africa, Canada (Nfld, Central Saanich)
What golden nematode does
burrow in to roots, feed on root tissues, cause death/stunted growth
Caenorhabditis elegans
model organism for developmental genetics- tiny (few mm’s), short generation times (3days), hermaphroditic (self-fertilize), eutely (959 somatic cells)
why C elegans is a model organism
cell lineage known for all cells - map
synaptic connections btw neurons mapped
entire genome sequenced
ratio of animals that are arthropods
4/5
Phylum Arthropoda importance
high successful by any metric important food web component medical importance economic importance model organisms
Arthropods success
most specious, most individuals, ability to invade almost all habitats, etc.
Arthropods in food webs
important component - 1º and 2º consumers, removal of arthropods would collapse any ecosystem
e.g. Arthropod medical importance
mosquitos are disease vector
e.g. Arthropod economic importance
positive and negative
pollination, crop destruction
Arthropod phylogenetic controversies
monophyletic or polyphyletic?
sister groups?
relations amongst major groups?
one of the most extensively debated subjects in evolutionary biology
Arthropod metamerism
convergent with Annelida
consolidations of multiple adjacent metameres into coherent morphological units with specialized functions
tagmosis
unit = tagmata
Arthropod exoskeleton
chitin (polysaccharide) + crosslinked proteins (sclerotization) secreted by epidermal epithelium
Arthropod exoskeleton functions
support
protection (predators, mechanical abrasion)
facilities movement
exoskeleton movement function
transmits force of muscle contraction
Parts of arthropod exoskeleton
epicuticle (lipids, waxes)
Protocuticle (exocuticle + endocuticle)
Epidermis
axial arthropod skeleton
4 hardened plates (sclerites): top = term, bottom = sternum, sides = pleuron
appendicular arthropod skeleton
articles - thin hollow tubes connected to form appendages
arthropod jointed appendages
joints formed by thin, flexible exoskeleton (articular membrane)
movement of arthropod appendages
antagonistic muscle bands
condyle
many appendages/articles
Arthropod antagonistic muscles
extensor and flexor muscles
condyle
one article fits precisely into the other - only permits movement in 1 direction
Benefit of many articles in arthropod appendages
movement in different directions/planes = large range of motion
Adaptive potential of arthropod appendages
versatile raw material:
- can be sculpted into diff. shapes/functions
- each article can be precisely moved by muscles
exoskeleton tools
sensory, mouthparts, prey capture, crawling, swimming, escape behaviour
arthropod secondary body compartment
hemocoel separated by diaphragm in to pericardial and peravisceral
Arthropod nephridium
epithelial tubes capped by epithelial sac = metanephridium
excretory tubes homologous to metanephridium
saccule homologous to shrunken coelom(?)
extra consequences of exoskeleton
moulting
sensilla
Arthropod growth
periodic moults under hormonal control- secrete inactive chitinases
moulting process, cellular
glandular cells secrete inactive chitinase (proenzyme) - new epicuticle secreted - proenzyme activated - digest endocuticle - split out at areas of weakness
new exoskeleton
week/soft, not cross linked - pump up with air/water -make bigger before hardening - harden - shrink back down - have room
sensilla
sensory, ball and socket joint, neurons w/ dendrites, lower exoskeleton cover
What is Arthropod sister group
Onychophora
Phylum Onychophora
velvet worm, low species #s compared w/ arthro. (180), very humid terrestrial, chitinous cuticle, metameric, non-jointed appendages, annelid/arthropod, distinct terminal ends (claws)
earliest known Arthropod
Hallucigenia, Burgess Shale, Field BC, early Cambrian
Arthropoda subphyla
Trilobitomorpha
Chelicerata
Mandibulata
Chelicerate tagmata
2 tagmata
Prosoma
Opisthosoma
Chelicerate # of appendages
Prosoma - 6 pr.
Opisthosoma - variable
Chelicerata first pr appendages
chelicerae
Chelicerata 2nd pair appendages
pedipalps
Chelicerata classes
Merostomata
Arachnida
Pycnogonida
Merostomata
1 genus (4spp.) extant, marine, horseshoe crab
Merostomata e.g.
Limulus sp.
horse shoe crab
Merostome morphology
2 tagmata: prosoma (A), opisthosoma (P);
telson, compound eye (unique)
Merostome prosoma
shovel-shaped to facilitate sediment burrowing
telson
not true metamere, hinged to body, d-v movements, aids in ‘righting’ flipped over body
Merostome appendages
Prosoma: chelicerae, pedipalp, (mouth), 4 walking legs
chelate
pincer-like appendage
article
podomere (segment of appendage)
Merostome posterior appendages
not chelate
‘pusher legs’
push off sediment
clean out gills
Merostome opisthosome
6pr appendages fused in to flaps = gill opercula
Merostome gill opercula
thicker exoskeleton protecting book gills (very thin exo.)
wave to oxygenate gills
gnathobase
proximal joint/ process of arthropod appendage, modified to aid in carrying/ masticating food
Class Arachnida
scorpions, spiders, ticks, mites
terrestrial, largest group of chelicerata, few opisthosome appendages, predators, malpighian tubules, book lungs/tracheae, spermatophores
Arachnid feeding
mostly predatory, mostly on other arachnids
How to feed on arachnids
must get in to exoskeleton - rip up, vomit digestive enzymes, suck up sap
malpighian tubules
lined tubules extend from gut and float in hemocoel, take up rates and crystallize- move down tubule - removed from anus - save water
Book lung
internalized book gill
delicate epidermis layers covered by thin exoskeleton
tracheal tubules
tubular, branched, invagination of epithelia w/ exoskeleton floating in hemocoel
Arachnid reproduction
internal fertilization
spermatophores - protected bundles of sperm
scorpion appendages
Prosoma: chelicerae (small), 1 pr pedipalps (large, chelate), 4 pr walking legs
telson: stinger w/ neurotoxin glands
Scorpion pedipalps
prey capture, ambush predator (only use stinger for big prey)
pectine
comb-like sensory structure - high density of mechanoreceptors to detect substrate movements, opens to book lung, ventral side of prosoma
Spider characteristics
no compound eyes, unique pedipalps, pedicel, spinneret
Spider pedipalps
fm - sensory appendages
m- deliver sperm
not chelate, daggers
Spider waist, between opisthosome and prosome
pedicel
narrow for movement = flexibility of spinneret
spinneret
many spigots release strands of silk
spider fertilization
no spermatophores, extrude sperm on to silk, pedipalp sucks it up in to chamber, fits in to fm gonopore like lock&key
functions of spider silk
sperm transfer egg cases prey capture aqualung ballooning
ballooning
move through the air by releasing threads to catch the wind
mites/ticks
huge diversity- aquatic, predator/ herbivorous/ parasites
mites/ticks appendages
hypostome
chelicera
pedipalp
4 pr walking legs
mites/ticks parasitic feeding
attach to host - slice open host w/ chelicera - insert hypostome to anchor and hold position - use pedipalps to support body and for sensory
Class Pycnogonida
sea spiders
ectoparasites, very small, puncture hole in host and suck out body fluid
Pycnogonid body parts
Prosoma: proboscis, chelicera (chelifore), pedipalp, ovigerous leg, 4 prs legs
Opisthosoma great reduced, bears anus
pycnogonid proboscis
forced through body wall of host to suck body fluid, chelicera help open wound for insertion
pycnogonid oviparous leg
often only in m, used to carry eggs, males carry and brood eggs until hatching
Arthropod groups
Chelicerata
Mandibulata
Mandibulata groups
Myriapoda
Pancrustacea
Pancrustacea
Crustaceans = Hexapoda
Ostracodes, Cirripedia, Malacostraca, Copepoda, Branchiopoda, Hexapoda, Remipedia
Is crustacea monophyletic
no, paraphyletic
mandibulata tagmata
2 or 3
head + trunk
or head+ thorax + abdomen
Mandibulate head + thorax
cephalothorax (secondarily consolidated)
tagmata in majority of Mandibulates
3
head+ thorax + abdomen
Mandibulate appendages
head: 1-antennae, 2-antennae or X, 3-mandibles, 4-maxillae I, 5- maxillae II
(3prs mouthpart appendages)
grasshopper mouthparts
disarticulated upper lip (labrum), 2 mandibles, 2 maxillae, lower lip (labium)
biramous arthropod appendages
epipod, protopod
biramous appendage protopod
2 basal articles made of multiple articles - exopod, endopod
compound eyes
common in mandibulate
many hexagons = ommatidium
ommatidium
single photoreceptor cartridge: each one samples section of visual field at different angle than neighbour, detecting slightly different visual field
after ommatidium detects image
crystalline cone focuses incoming light - reticular cells covered in microvilli are receptors - open ion channels - membrane polarization - propagate action potential - brain -information
Myriapoda tagmata
2:
head
thorax
Myriapoda appendages
head: pair antennae, mandible, 2 pr maxillae
What are myriapods
centipedes, millipedes
centipedes
predators, poison claws, fast
centipede poison claw
first trunk appendage
centipede movements
fast effective stroke, slow recovery stroke, high speed, high gear, low power, not many legs on ground at once
Millipede
herbivorous, slow, diplosegments
diplosegment
2prs legs/segment
millipede movement
slow effective stroke, fast recovery stroke, slow movement, low gear, high strength, most legs on ground at once pushing - strong walking
What is hexapoda sister group
Remipedia - primitive body plan - annelid like, not very successful
Hexapoda terrestrial adaptations
Minimize water loss
Avoid/tolerate temperature extremes
Wings/flight
Hexapoda water loss
minimized via: waxy epicuticle (myriapods lack waxes in epicuticle), mapighian tubules, water reabsorption (specialized rectum), tracheal tubules
Hexapod malpighian tubules
tubules in hemocoel take up urates, precipitate uric acid (non-soluble), pass w/ feces
glow worm
glow produced w/ malpighian tubules - sticky silk threads ensnare prey
Hexapod tracheal tubules
hollow cylindrical invaginations of body wall, dendritically branched, end on cell body, O2 delivered directly to cell via tube, lined inside by exoskeleton
opening of hexapod tracheal tubule
spiracle with filter hairs
tracheal tubule support
Taenidia - circular thickenings of exoskeleton along length of tube to prevent collapsing
Myriapod spiracles
not closable (dry out quicker)
Hexapod temperature tolerance
behavioural
morphological
physiological
behavioural temperature tolerance, hexapod
basking, stilting, crouching
stilting
extend legs, lift body off of hot ground
crouching
ground heats/cools slower than air, crouching close to rock to absorb heat
morphological temperature tolerance strategies, hexapoda
insolating cuticler hairs, layer of hair around body prevents heat loss, e.g. bumblebee
physiological temperature tolerance strategies, hexapoda
shivering, antifreeze proteins
dangers of freezing
sharp crystals rupture cells
AFP
Antifreeze Proteins
Insect AFP
surround small ice crystals and prevent them from growing
Hexapoda wings
mesothoracic, metathoracic wings
from 2nd, 3rd thoracic segments
wingless insects
apterygotes
insect wing morphology
largely exoskeleton, reinforced by wing veins (tracheal tubules), nerves, sensory receptors, tissues
Advantage of flight
new niche space, dispersal to new resources, reach resources inaccessible to others, escape predators, find mates, migration, find suitable mating locations
origin of wing hypotheses
Tergal lobes
Appendage derivatives
Tergal lobe theory
tergum (top sclerite lobe) draw out laterally
problems with tergal lobe theory
no hinge (wings are hinged) would require elaborate modification
Appendage derivative hypothesis
epipods are hinged and fn in gas exchange
testing appendage derivative hypothesis
Drosophila gene rubbin (Tc factor) is essential for wing development; mutated rubbin = little nub wings; look for homologues of rubbin in pancrustacea - mark - follow development
results of appendage derivative hypothesis testing
rubbin traced to epipod in crayfish and brine shrimp- consistent with epipod-wing theory
Hexapoda mouthpart diversification
chewing, sponging, piercing, sucking
ancestral hexapod mouthparts
chewing type
e.g. grasshopper
grasshopper mouthparts
heavily sclerotized, chewing:
labrum, 2 mandibles, 2 maxilla, labium (2nd pr maxillae), hypopharynx
winged insects
pterygotes
hypopharynx
tongue
salivary glands
sponging mouthparts, hexapoda
labium (2nd pair maxillae)
tubules for sponging liquid food
cutting + sponging mouthparts
sponging labium + cutting mandibles (cut open prey to sponge up fluids)
piercing and sucking mouthparts
labium =protective sheeth, supports stiletto-like mouthparts - labrum, mandibles, maxillae = elongate spheres;
mandibles puncture, labrum forms sucking tube, salivary secretions make us itchy
butterfly mouthparts
sucking; 1st pr maxillae curve together to form elongate sucking straw
Hexapod development patterns
ametabolous
hemimetabolous
holometabolous
exopterygote development
=external wing development
hemimetabolous development
larva similar to adult minus wings
Ametabolous development
apterygotes
nymphs hatch from eggs that look exactly like adult, live in same habitat, feed on same material
stages of moults all look same
no major transitions in development (except reproduction)
how are hemi/holometabolous youth different than adults
mainly they do not have wings
holometabolous development
endopterygote development
larva very different than adult
internal wing development
example of hemimetabolous development
grasshopper, locusts, mayfly larva, dragonfly larva
what do hemimetabolous wings develop from
wing pads
what do holometabolous wings develop from
imaginal discs
holometabolous specialties
differential specialization
larva specialized for feeding
adult specialized for reproduction
holometabolous metamorphosis
body rearrangement
exclusively adult features develop internally (wing, antennae, mouthparts)
imaginal discs
nests of stem cells that differentiate in to various adult structures
pupa stage, holometabolous development
rearrangement
larval structures destroyed
adult structures everted
aquatic larva, Hexapoda
2 life history stages, major development, differential specialization
mosquito life cycle
adult - eggs in water - larva (w) - pupa (w) - adult emerges in to air
larva has posterior siphon for breathing
pupae has anterior snorkle
Hexapoda defensive strategies
morphological
behavioural
chemical
physiological
Hexapod morphological defensive stratgies
shape and colour matching
e.g. stick bug
hexapod behavioural defensive strategies
flight
projectile defecation
petiole clipping
behavioural crypsis
projectile defication
release feces in projectile motion to hide location
petiole clipping
fill up on leaf, chop off stem so hole-y leaf is not so visible
example of behavioural crypsis
woolly aphid, ant, lacewing larvae
example 2 of behavioural crypsis in hexapoda
leaf beetle larva
hold mass of own feces containing defensive chemicals from plant on back - disguise as something not tasty
chemical defensive strategies, hexapoda
venomous stings (bees, wings)
sticky threads to entangle
(termites)
reflex bleeding (ladybug)
relex bleeding
toxic hemal fluid released from self-directed rupturing of articular membrane between leg articles
articular membrane
thin area of exoskeleton ‘joints’
bombardier beetle chemical defence
abdominal reaction chamber - explosively eject hot, toxic fluid from anus
bombardier beetle reaction
secrete hydroquinone and H2O2, catalaze reduces H2O2, O2 oxidizes hydroquinones (exothermic), hot quinones released (toxic)
Non-hexapod pancrustacean characteristics
2 pair antennae (hexapods have 1)
biramous appendages (hexapods have uniramous)
nauplius larvae
Nauplius larva
no cilia
3 pr muscle-operated appendages
single median eye
2 pr antennae, 1 pr mandibles = swimming
Pancrustacea clades we focus on
Copepoda
Cirripedia
Malacostraca
Hexapoda
generalized Malacostracan body plan
3 tagmata - abdomen (6 segments), thorax (8), head (5)
Malacostracan head appendages
2pr antennae, 1 pr mandibles, 2 pr maxillae
Malacostracan thorax appendages
thoracopods
8 prs appendages
maxillipeds, pereopods
Malacostracan maxillipeds
1+ pairs often modified accessory mouthparts
Malacostracan pereopods
walking legs
Malacostraca abdomen appendages
6pairs
pleopods (swimming)
uropods
Malacostracan head/thorax consolidation
tergal sclerites fused in to cephalothorax = carapace
lateral carapace flaps form chamber
carapace gill chambers, Malacostracans
branchiostegites
Euphausiacea
Malacostraca, krill
incomplete branchiostegite, no maxillipeds, specialized thoracopods
Euphausiid thoracopods
setose biramous feeding appendages = feeding basket/sieve
setose
exoskeleton elaborated in to bristle like extensions
Pericarida, Malacostraca
Amphipoda, Isopoda
no carapace, 1pr thoracopods = maxilliped, eggs brooded in marsupium formed by oostegite
Amphipoda
laterally compressed
Isopoda
dorso-ventrally compressed (form ball)
coxa
basal most article
basal
bottom layer, closest to body
Caprellid amphipod
skeleton shrimp, Malacostraca
stomatopod
mantis shrimp (Malacostraca) tropical/semitropical, benthic, carapace does not cover entire length of thorax, functional telson, very aggressive
Stomatopod appendages
5prs maxillipeds, 2pr raptorial appendages
spearing appendages, stomatopod
species that burrow in soft substrate
smashing appendages, stomatopods
species that live in rock crevices
“most complex visual organ on the planet”
stomatopod compound eye
Stomatopod eye
mounted on moveable stocks = 360ºrotation = large visual field
eye divided in half = stereo vision = broad focal range
12-19 photoreceptors w/ different photopigments
human photoreceptors
3
what is the different in having more photoreceptors if they cover the same spectral range
more sensitive to subtle difference in colour, and extremes (UV, polarized)
less ‘windows’ (missing coverage areas)
Stomatopod telson functions
wide flat defense shield
reflect polarized light -signaling?
Decapoda groups
Caridoa
Astacidea
Anomura
Brachyura
Decapod general characteristics
(Malacostraca)
well-developed carapace (entire thorax)
3 pr maxillipeds
Caridea
Shrimp delicate, slender pereopods large muscular abdomen well developed pleopods holopelagic or pelago-benthic
Caridea pereopods
delicate perching appendages
unique amongst the decapods
Caridea pleopods
well developed, swimming
Astacidea
Crayfish, lobsters (Decapoda, Malacostraca)
massive pereiopods, large muscular abdomen, pleopods, chelipeds
Astacidea pleopods
swimming (but don’t swim a lot)
Astacidea chelipeds
1st pair of pereopods - prey capture, defense, offence
Brachyura
true crabs (Decapoda, Malocastraca) broad flat cephalothorax, heavy robust pereopods, chelipeds, reduced tucked in abdomen, closed gill chambers
Brachyura appendages
robust pereopods 1st pr pereopods = chelipeds uropods - secondarily lost pleopods - reduced/lost, retained in fm to hold eggs between thorax and abdomen 4 pr walking legs
Anomura
squat lobsters, hermit crabs (Decapoda, Malacostraca) most diverse group of decapods very reduced 5th pr pereopods 3 prs walking legs abdomen well developed
hermit crab
abdomen has spiral asymmetry to fit gastropod shell
pleopods only developed on one side
uropods have grippers to hold on to columella
Porcelain crab
Anomuran
carcinization
3 prs walking legs
carcinization
anomurans that have converged body morphology with brachyuran crabs (reduced abdomen, carapace over cephalothorax)
gill bailer
decapod elaboration of second pair of maxillae, aerate gills, moves in a wave-like fashion
why do decapods need gill bailer
to clean out gill filaments - no motile cilia!!
Caridea gill cleaning
chelate pereopods - reach in to gill chambers w/ delicate pinchers
(no chelopeds)
Anomuran gill cleaning
5th pair of pereopods = little ‘stub’ on posterior dorsal end of carapace, insert in to gill chamber for cleaning, setose tip
Brachyuran gill cleaning
3pr maxilliped epipods = elongate, setose, gill cleaning combs
extend back in to gill chambers
gill scrubbing whenever maxillipeds move
Brachyuran, carbonized Anomurans
closed gilled chambers
lateral rim of branchiostagites fused to abdominal surface
Anomuran tail fan
telson + 2pr biramous uropods
used along w/ abdominal muscles for movement = tail flip
Tail flip, jump backwards
flexion of posterior abdomen segments - pulls body backwards
Copepoda lifestyle, habitat
Pancrustacea
extremely abundant, diverse aquatic habitats - freshwater/marine- puddles, hot springs, enormous number of individuals, largest animal biomass on the planet, holopelagic, pelago-benthic, parasitic
Copepod body size
1-2mm
Copepod importance
transfer organic carbon from producers to higher trophic levels
tail flip, jump up
flexion between abdomen and cephalothorax -bend middle of body – body move upwards
Copepod body
torpedo-shaped
2 tagmata - head/thorax, abdomen
single medial eye
copepod appendages
1st pr antennae - sensory 2nd pr antennae- swimming pr mandibles, 1st pr maxillae 2nd pr maxillae - food capture 1pr maxillipeds - assist feeding 4-5prs thoracic app. - hop swim forked telson - 2 caudal rami
distinctive copepod features
forked telson
copepod swimming appendages
2nd pr antennae (biramous, bristled)
4-5prs thoracic appendage (hop swim) coupled together along median line
copepod feeding appendages
2nd pr maxillae
1 pr maxillipeds
Planktonic copepod environmental factors
pull of gravity depth-dependent light intensity nowhere to hide dilute resources small Re #
Maintaining position in water column, copepods
1st pr antennae long,bristled increase drag
2nd pr antennae- swim
store nutrients as lipid globules for buoyancy
Reynolds number
describes viscous : inertial forces for characterizing behaviour of fluids flowing past an object
Re =
(velocity x size x density) / viscosity
Copepod feeding
flow lines disrupted by particle - 1st antennae detect disrupted flow while swimming on back- 2nd pr maxillae move apart rapidly - negative pressure, particle drawn in - close maxillae rapidly to capture particle
Re dominated by
velocity, size
density/viscosity relatively unchanging within water column
Re less than 10
flow lines move past object in orderly fashion, maintain trajectory, at low speed fluid dominated by viscous forces
Re above 20,000
Turbulent flow
dominated by inertial forces
flow lines severely disrupted past object
copepod swimming
very active movement, no gliding when movement stops
small size = low Re = viscosity dominant
Why do copepods move maxillae rapidly to feed
to increase Re (increased velocity)
boundary layer
non-moving fluid around object
low Re = thick boundary layer
flow dominated by viscous forces
small Re
how do copepod detect changes in flow lines
1st antennae covered in mechanoreceptors and setose bristles have thick boundary layer (paddle-like)
Copepod evasion and escape
transparent w/ few tissue pigments loss of compound eyes (pigmented) hop swim diel vertical migration 1st antennae sensory axons have myelin sheath
copepod myelin sheath antennae
action potentials travel more rapidly if axon is enlarged (vertebrates), inverts use myelin sheath to speed potential
Copepod mate finding
Pheromones
Behaviour patterns
how copepod mating strategies work
male swim back and forth across top of column, fm swim up and down releasing pheromone, when pheromone detected m swims down
thin boundary layer
large Re
copepod reproduction
eggs brooded in egg sacs
nauplius larva
parasitic copepod example
salmon louse
ectoparasite
huge, cling to external surface, feed on mucus/epidermis/ blood, detrimental to fish health
giant parasitic copepod
family Pennellidae
ectoparasite of marine mammals
up to 30cm long
Class Cirripedia
sessile in post-metamorphic stage (unique)
carapace as calcified plates
suspension feeding
free-living or symbiotic
main Cirripedia groups
acorn barnacles
stalked (gooseneck) barnacles
acorn barnacle
calcified cone directly attached to substrate
stalked barnacle
calcified plate mounted on fleshy stalk
Cirripedia morphology
opercular plates
wall plates
cirri
Cirri
6 pairs thoracic appendages, biramous and setose, very small = low Re, feeding paddles
flow dominated by inertial forces
Large Re
generate turbulent flow
Gooseneck barnacle environment
high flow -incapable of sweeping cirri, can’t raise Re
passive feeding
Gooseneck barnacle morphology
peduncle (stalk)
adhesive gland
ovary, gut, cecum, muscle, mouth, mantle cavity, cirri
Cirripedia moulting
partial moult, only exoskeleton of cirri
Cirripedia growth
CaCO3 added to basal rim and up sides - grow in diameter, height
barnacle reproduction
most hermaphroditic
very long penis
gregarious settlement, metamorphosis
gregarious settlement
larval stage attracted to settle near other members of the species - need to be nearby each other
benefit of hermaphroditism in barnacles
as long as there is neighbour fertilization can take place
barnacle life cycle
Nauplius larvae - cyprid larva
Cyprid larva
2nd larval stage, non-feeding, must find suitable settling place, crawl around on rocks w/ 1st pr antennae
symbiotic barnacles
commensal
parasitic
commensal barnacles
dwarf, complemental males in some species
when density low
settle out of copulation range
barnacle on crab under anemone
decorator crab with anemone on top to protect itself against cephalopods - barnacle reaches up and rips off anemone tentacles
Rhizocephalan barnacle
endoparasite of decapod malacostracan
highly derived adult morphology
manipulation of hosts behaviour
Rhizocephalan life cycle
dioecious - naplius metamorphoses to cyprid - fm swim around looking for decapod - settle - slice hole- insert cells - grow in to series of branched root-like structures that invade all tissues of host (interna) -derive nutrients from host - break through as mass of tissue to exterior (externa) - ‘mate’
Rhizocephalan mating
fm forms externa, male cyprid finds externa, metamorphosis into dwarf male - essentially a sperm sac
where does externa go
breaks out where egg mass would normally be, alters hosts behaviour to treat it like eggs (aerate, clean) - even male hosts!
Phylum Echinodermata major characteristics
'spiny skin' Deuterostome Eucoelomate WVS calcareous endoskeleton pentamerous radial symmetry mutable collagenous tissue
Echinoderm body compartments
coelom
endoderm derived mesoderm
3 sets of compartments formed in development
Echinoderm coelom development
enterocoely
WVS
water vascular system
coelomic compartments
operates tube feet
tube feet
highly flexible, muscular, tubes filled d w/ fluid, moved by hydrostatic mechanism, unique
pentamerous radial symmetry
pentaradial
at least in adult stage
bilateral + 5 point
bilateral in juvenile stage
mutable collagenous tissue
connective tissues that can change response to tension between extensible and rigid, under control of nervous system
Echinoderm skeleton
calcareous endoskeleton spines are skeleton covered w/ living tissue secreted by embryonic mesoderm ossicles microporosity
do all echinoderms have mutable collagenous tissue
yes, but not all collagenous tissues are mutable
normal connective tissues
lots of collagen fibres
form in keeping things together (skin to muscle) - generally not active
Echinoderm subphylum
Crinozoa
Asterozoa
Echinozoa
Crinozoa groups
Class Crinoidea
Asterozoa groups
Class Stelleroidea - Subclasses Asteroidea, Ophiuroidea
Echinozoa groups
Classes Echinoidea, Holothuroidea
Asteroidea
sea stars, mostly predatory
spines, disk, ambulacra, madreporite, arms, ossicles, pedicellariae
ambulacra
where tube feet extend to environment (oral surface)
Asteroid endoskeleton
latticework of interconnected ossicles, laced together w/ collagen fibres
spines
Asteroid pedicellariae
2 jaws formed by specialized ossicles
cleaning, removing settlers
Pedicellariae morphology
epidermis
2 jaw ossicles
basal ossicle
attached by opener/closer muscles
Echinoderm canals
fluid-filled tubes ring canal around esophagus/ mouth radial canal down arm lateral canals out from radial to tube feet stone canal - from ring to madreporite
movement of tube foot
close valve - contact muscle surrounding ampulla- force water down in to tube foot- foot extended - reach out- contract tube foot muscles to bend
tube feet are like
flexible pipettes
polian vessicle
filled with fluid
fluid reservoir
stone canal
reaches up to specialized ossicle (madreporite)
madreporite
very coarse ossicle
replenish fluid in WVS by taking in sea water
Tiedmann’s body
manufactures phagocytes to phagocitize invading particles that come in madreporite
Echinoderm coelomic compartments
WVS
perivisceral coelom
perihaemal coelom
genital coelom
Echinoderm nervous system
ring nerve + 5 radial nerves
intra-epithelial
no centralization
same organization as WVS
sea star feeding
tube feet pry open shell- evert cardiac stomach- digest and ingest
specialized echinoderm ossicles
madreporite
spines
pedicellariae
Asteroid reproduction
Asexual - fission and regeneration, autotomy
Sexual
planktonic, feeding larva
catastrophic metamorphosis
Asteroid fission
central disk breaks in two then regenerates missing parts
Asteroid autotomy
arm is shed and lives independently as a ‘comet’, eventually regenerating missing parts
Asteroid sexual reproduction
dioecious
broadcast spawn
external fertilization
respiratory out foldings of sea star body
papulae
‘no arm’ Echinoderms
Echinozoa (Echinoidea, Holothuroidea)
Echionoidea
no arms
endoskeleton
aristotles lantern
echinoidea endoskeleton
flat ossicles that fit tightly together
5 ambulacral plates, 5 interambulacral areas, 2 rows of plates in each
spines, stalked pedicellaria
ambulacral plates, echinoidea
2 rows of double pores for tube feet
sea urchin spines
attach via ball-and-socket joints, moved by muscles, function in bracing, manipulating food, defense, may extrude toxin
Echinoid pedicellaria
stalked
calcareous support rods
three opposing jaws
Aristotles Lantern
complex system of ossicles and muscles surrounding esophagus
teeth can be protruded from mouth and moved in various directions to eat or scrape
regular echinoids
sea urchin
radial symmetry
rocky substrates
irregular echinoid symmetry
superimposed bilateral symmetry
e.g. sand dollar
regular echinoid locomotion
spines and tube feet
regular echinoid feeding
scarpe algae from rocks
shred kelp
capture drift algae with tube feet
Irregular echinoids
sand/mud substrate
locomotion by spines only
deposit feeding (tube feet)
Irregular echinoid respiration
petaloid tube feet specialized for gas exchange
Holothuroidea body
no arms
elongation along oral-aboral axis
bilateral symmetry
Holothuroidea endoskeleton
microscopic ossicles
not connected flexible, muscular body wall
Holothuroidea WVS
5 ambulacra
buccal podia
suspension/deposit feeding
internal madreporites
Holothuroid ambulacra
bivium (‘dorsal’)
trivium (‘ventral’)
buccal podia
circle of 10-30 tentacles around mouth, may be same
size, or some dwarfed, may be branched (dendritic / arborescent), may be pinnate, peltate or digitate, retractile and the body wall can close over them
cuvierian tubules
toxic structures attached to left respiratory tree, used for discouraging and entangling potential predators
evisceration
expelling internal organs, cuvierian tubules or entire digestive system, respiratory trees, and gonads; liquify and rupture connective tissue attaching viscera to inner body wall, eventually regenerated
Holothuroidea distinctive characteristics
sausage shape microscopic ossicles muscular body wall spacious perivisceeral coelom buccal podia for feeding respiratory trees
ophiuroid suspension feeding
bury central disk and stick arms up (kind of looks like whip coral)
Basket star
ophiuroid w/ branched arms - suspension feeding, coil around prey (zooplankton)
bursal slit
narrow slit along inner arms of Ophiuroids, leading into large thin-walled sac (the bursa) which has a respiratory function
Ophiuroidea characteristics
central disk w/ highly flexible arms locomotion by arm rowing arm autotomy (MCT) no anus tube feet lack suckers and ampullae bursa for gas exchange
MCT
mutable collagenous tissue
soften body wall before autotomy
brittle star respiration
cilia-lined sacs called bursae; each opens between the arm bases on the underside of the disk
Crinoidea
‘sea lily’, ‘feather star’
arms with pinnules and podia
stalk or cirri
Crinoidea feeding
suspension feeding w/ pinnule podia
Crinoid endoskeleton
disc or vertebrate-shaped ossicle
allow long term extension of arms
crinoids as representatives of primitive echinoderm state
body orientation - mouth up
endoskeleton -articulating discs
WVS for suspension feeding (not locomotion)
MCT for long-term maintenance of posture w/ no E expenditure
Deuterostomia phylogeny
[Echinodermata s.g. Hemichordata]
s.g.
Chordata (Cephalochordate s.g. Urochordata s.g. Craniate)
Chordates
perforated pharynx
notochord
dorsal hollow nerve cord
muscular post-anal tail
Subphylum Urochordata groups
Class Ascidiacea
Class Larvacea
Class Thaliacea
Urochordata characteristcs
also Tunicata
Tunic
no metamerism
eucoelom as pericardium
tunic
tunicin cellulose-like fibrous material
Class Ascidiacea
sea squirts
sessile suspension feeders
solitary and colonial forms
Ascidian tunic
epidermis secreted tunicin + protein
hemal channels w/ wandering blood cells
spicules, fibrous material
branchial basket
cartilaginous structure supporting the gills in protochordates and lower vertebrates
buccal siphon, atrial siphon
w/ pharyngeal mucus net to catch particles that go through perforated pharynx
colonial ascidians
zooids interconnected by stolons
zooids rise from basal mat of shared tunic
zooids entirely embedded in shared tunic
Didemnum sp.
invasive species of colonial tunicate, aggressive space competitior
Ecteinascidia turbinata
Ascidian, source of anti-cancer drug ‘yondelis’ - PharmaMar, now synthetically manufactured
Ascidian reproduction
hermaphroditic
broadcast spawn
eggs w/ self/non-self recognition
tadpole larva
Ascidian larva
tadpole short-lived, non-feeding tail w/ notochord, dorsal hollow nerve chord, muscles adhesive papillae start of branchial basket
Ascidian metamorphosis
tadpole attaches to substrate w/ adhesive papillae (anterior end)
retraction of tail (apoptosis)
90º rotation of viscera
viscera
internal organs in the main cavities of the body, especially those in the abdomen
Class Larvacea
planktonic, 1-2mm
disposable gelatinous house
pharynx w/ 2 stigmata, mouth, anus, gut all in ‘head’
tail w/ notochord and dorsal hollow nerve chord
larvacean house
secreted by epidermal epithelium (4-8/day) blown up by tail thrashing captures food particles escape opening pre-filters food-concentrating filters animal (mouth, tail) excurrent opening
Heterochrony
evolutionary change in time of appearance or rate of development of a character relative to other characters
e.g. somatic tissues vs reproductive tissues
Heterochrony example
somatic tissue arrested development, reproductive tissue development = sexually mature ‘juvenile’
possibly why larvacean looks similar to Ascidian tadpole
Heterochrony results
paedomorphosis
peramorphosis
paedomorphosis
adult of a descendant is similar in appearance to the juvenile (larva) of its ancestor
Class Thaliacea
pelagic, colonial (at least some stages)
pharyngeal cilia for swimming and feeding
individuals resemble adults ascidians
Thaliacea colony
pyrosomes, up to 10+m long
hollow pelagic tubes, closed at one end, walls contain many zooids
communal tunic
bioluminescent
individual Thaliacea zooid
perforated pharynx, siphons, like mini ascidian
exhaling water released into lumen of colony - expelled out colony aperture - propels colony along
thaliacea colony communication
no interconnected neurons but coordinated movements
chain rxn - organism hits something - flashes- neighbour detects w/ photoreceptor - flashes - propagate message
Phylum Hemichordata group
Class Enteropneusta
Also Class Pterobranchia but not for our purposes
Class Enteropneusta
Acorn worm (anterior looks like acorn)
live in soft sediment
secrete a lot of mucus
eucoelomate
Enteropneusta body
tripartite: proboscis, collar, trunk
tricoelomic (enterocoely similar to echinoderm)
Enteropneust burrowing
peristaltic muscular contractions of proboscis
proboscis w/ cilia, mucus - pick up particles and carry posteriorly to move sed out of way - start burrowing motion- muscularly continue burrowing
Enteropneust feeding
crawl up burrow to surface, lay proboscis on surface, cilia ensnare particles, carry to mouth
back up to surface to release fecal casting
Acorn worm respiration
2 sets of perforations in external body wall = gill pores, perforated pharynx in dorsal region
ingested water travels out branchial pores and then external gill pores
Acorn worm nervous system
D and V intraepithelial nerve cords (like sea star)
intra-epithelial nerve plexus (nerve tails run out to all parts of body)
hollow dorsal nerve cord in collar
Enteropneust movement due to
ciliary activity and muscular contractions
juvenile (larva) of a descendant is similar to the adult of its ancestor
Peramorphosis
Acorn worm dorsal hollow nerve cord?
short region in collar
intraepithelial nerve cord sinks down and pinches off to become a d.h.n.c.
Enteropneust reproduction
dioecious
larva morphologically similar to holothurian
broadcast spawn
ciliated larval stage may feed for months
Hemichordate echinoderm-like characters
larval morphology (similar to some groups, holothurians) intraepithelial nerve system (found in sea stars and trunk region of hemichordates)
Hemichordate chordate-like characters
pharyngeal perforations
dorsal hollow nerve cord (collar)
Ctenophora
mostly planktonic, swim but not strong enough to resist current all marine and carnivorous 100-150spp., but large individual #'s mm - 1.5m often translucent
Ctenophora body
gelatinous - highly hydrated mesoglea (similar to cnidaria)
tentacles
comb rows/plates
mouth up - pharynx - stomach - anal pores
gastrovascular canals
Ctenophore tentacles
2, originate from pits (pentacular sheaths), can be retracted, have side branches (tentilla) loaded w/ collocytes
Ctenophore comb plates
8 rows of comb plates (ctenes), used for swimming, extend from oral surface down
ctenes
large ciliary structures known
thousands of cilia in a row = plate
move in unison, out of stroke down comb row - wave propagates down comb row
cilia connected by interciliary links
ctenophore cilia
up to 2mm
largest animals that locomotive by cilia
ctenophore movement
effective stroke is towards aboral end
move through water column with mouth leading
beat frequency can slow and switch directions
Collocytes
release sticky adhesive single use only differentiate from interstitial cells complex, used to capture prey head covered in intercellular granules straight filament = elongate nucleus
Ctenophore muscles
myoepithelial cells in epidermis
bona fide muscle cells within ‘mesoglea’
presence of stand-alone muscle cells in ctenophores
MAY suggest that ctenophores are true mesodermal animals and therefore triploblastic (no evidence)
ctenophore nervous system
nerve net in epidermis and mesoglea
aboral (apical) sensory organ (rudimentary brain)
ctenophore aboral sensory organ
mostly statocyst (gravity sensor) transparent CaCO3 dome over statolith on 4 balancers
statolith balancers (ctenophore)
bundles of hundreds of cilia
tracks bifurcate away from dome
how ctenophore apical sensory organ works
turn - statolith moves - pressure on 1+ balancer = signal down comb row = more rapid beating of cilia to make that side move up and straighten out
detect direction of gravity to know which way is up
Pleurobrachia feeding
prey captured w/ sticky collocates on tentilla
tentacle retracts
ctenophore spins to waft tentacle over mouth
Ctenophore defense
possibly cnidocyte stealer
feed on hydromedusa tentacles, place cnidocytes in ctenophore tentacles (observed in rare species)
Cestum spp.
Venus’ Girdle
long, thin, flat, ctenophore
Mnemiopsis
lobate ctenophore
invasive species, invaded black sea, no natural predator, more weight than annual world fish catch
capture prey w/ muscular oral lobes
Beroë
‘swimming mouth’, no tentacles, feeds on other ctenophores, introduced to bring down population of mnemiopsis
Beroë teeth
macrociliary teeth around margins of mouth, prevent prey from escaping, formed of cilia
Benthic ctenophores
look like flatworms
lack comb rows in adult stage
ctenophore reproduction
asexual - benthic ctenophore only
sexual - simultaneous hermaphrodites
-gametes from gastrodermis
-cydippid larva (look like pleurobrachia)
Bryozoan main characteristics
sessile, colonial, lophophore, defense strategies, polymorphic colonies
example of bryozoan heterozoid
Avicularium
Molluscan phylogeny
Conchifera (Cephalopoda, Scaphopoda, Bivalvia, Gastropoda)
Acquifera (Polyplacophora, Chaetodermomorpha, Neomeniomorpha)
largest class of Molluscs
Gastropoda
type of shell with mantle cavity over head
endogastric shell
representative of torsion
shell perforations =
2 ctenidia (left, right)
largest gastropod group
Caenogastropoda (predatory, use proboscis)
Cone snail toxin gland homologous to
mid-esophageal gland in other gastropods
bivalve ctenidia modification
adapted for suspension feeding
shell with mantle cavity at posterior end
exogastric shell
Nematode secondary body compartment
pseudocoel
Arthropod subphyla
Chelicerata
Mandibulata
Chelicerata groups
Merostomata (horseshoe crabs)
Arachnida
Pycnogonida (sea spiders)
Mandibulata groups
Pancrustacea
Myriapoda (centipede, millipede)
Pancrustacea groups
Hexapoda
Malacostraca
Copepoda
Cirripedia
Malacostraca groups
Euphausiacea
Stomatopoda
Pericardia
Decapoda
Decapoda groups
Caridea
Astacidea
Anomura
Brachyura
Merostomata appendages
1st appendages = chelicera, feeding
2nd appendages = pedipalps, variety of fn, not very derived
Insect tagmata
head
thorax
abdomen
Holometabolous development
complete reorganization of body
‘Whip’ heterozooid
Vibraculum