Marine Biology 278

Apr. 5, 2009 REVIEW OUTLINE FOR EXAM II

NOTE: you might want to review topics from EXAM I / REVIEW OUTLINE I that apply to the current material. E.g.: temperature and life; wave action; light colors & UV

BENTHIC BIOLOGY

LECTURES 14-15: OVERVIEW of BENTHIC ANIMALS: PHYLA as follows:

a. PORIFERA--filterers: pores (and exit osculum) and flagellated cells

b. CNIDARIA--carnivores: nematocysts, tentacles!! Polyp forms: Hydrozoa (simple,) Anthozoa (complex)

d. Platyhelminthes--carnivores: protrusible pharynx, hard tip stylet

e Pseudocoelomates: rotifers, gastrotrichs, etc.—MEOIFAUNA! (know definition)

f. Brachiopoda--filterer (spiral lophophore); clam-like 2-piece shell but has pedicle stalk & hole!

g. Bryozoa--filterer (tentacled /ciliated lophophore, horseshoe shape); zooids more advanced than polyps

h. MOLLUSCA—ARCHETYPE has head/foot; radula, shell. Features, feeding of classes:

i) Cl. Polyplacophora--8-plate shell; herbivores

ii) Cl. Gastropoda--1-piece shell; gone in slugs/nudibranchs; many feeding types

iii) Cl. Bivalvia--most filterers using gills; 2-piece hinged shell

iv) Cl. Cephalopoda--radula + beak: carnivores; shelled or reduced or lost shell (octopods)

i. Sipunculida--deposit feeder; sticky tentacles

j. Echiuroidea--filterers; proboscis, U-tube

k. Nemertina—ribbon worms, protusible sticky proboscis wraps prey

l. NEMATODA--many types of feeding; jaws; meiofaunal

n. ANNELIDA; Cl. POLYCHAETA--segments/parapodia, chaetes; many feeding types

o. ARTHROPODA --SubPHYLUM CHELICERATA: Cl Pycnogonida: seaspiders

p. ARTHROPODA subphylum CRUSTACEA; 2 pair antenna at front

i) Cl. Cirripedia--filterers (limbs); head grows into mantle=carapace/cement glands

iii) Cl. Malacostraca-- lobster, crab, hermit crab, many shrimp; isopods, amphipods

q. ECHINODERMATA--features of 5x-fold symmetry; skeleton; spines; tube feet

i) Cl. Asteroidea--carnivores (stomach evert) ii) Cl. Echinoidea--sand dollar, sea urchin types; Aristotle’s lantern; many feeding types; iii) Cl. Ophiuroidea--deposit feeders

iv) Cl. Holothuroidea--deposit, filter feeders; v) Cl. Crinoidea--sea lilies; mostly filter-feed

r. CHORDATA subphylum UROCHORDATA--2-siphon filtering system: Cl. Ascidea=tunicate/sea squirt

s, CHORDATA subphylum VERTEBRATA: many feeding types: i) Agnatha;

ii) Cl. Chondrichthyes iii) Cl. Osteichthyes v) Cl. Reptilia incl Cl. Aves vi) Cl. Mammalia

A. INTERTIDAL Zone

TECTONICS, Surface, Movements:: Basics of Rocky, Sandy, Muddy habitats and wave energy

TECTONICS: Active vs Passive margins -- differences and why

TIDES: SOLAR Tides create neap, spring tides: how

ROCKY SHORE COMMUNITY: Highest wave energy

ADAPTATIONS to Tide/Wave factors:

1. Wave Shock--pounding physically by waves, altered by tide level

a) Armor: shells ; b) streamlining ; c) elastic dissipation; Flexible polysaccharides in Macroalgae!

d) GLUES & anchorages: --mussel byssal threads!!

e) Behavioral Avoidance—crevices; “boring” clams

f) Life-cycle: i) colonies give mutual protection; ii) planktonic larvae to recolonize destroyed area

2. Water availability/loss (evaporation problem):

a) Armor: shell, other waterproof coverings

b) Physiological "Tolerance"--algae dry up 60-90%; some chitons 30-70%; unknown how survive,

c) Behavioral Avoidance--crevices/under algae

d) Life-cycle: i) colonies give mutual protection; ii) planktonic larvae to recolonize destroyed area

3. Oxygen: gills don't work in air

a) Lungs—Littorina periwinkle! Isopod scavengers -- both live in upper intertidal

b) Physiological Tolerance: Dormancy & enhanced Anaerobic metabolism:

i) non-acid endproducts (cf. lactic); or ii) branch to modified Krebs-cycle path; yield more GTPs/ATPs

4. Temperature -- Pattern: rapid ∆T in sun exposed limpets, barnacles, etc.

a) Coloration, shape: Reflect/radiate: light-colored and/or sculpted shells; darker to absorb in polar

b) Physiological Tolerance: temperature-insensitive dormancy; how stress or heat-shock proteins might protect upper intertidal organisms

c) Behavioral Avoidance--crevices/under algae

d) Life-cycle: planktonic larvae to recolonize

5. Salinity--drastic changes possible in air: osmosis damage

a) Armor--waterproof coverings

b) Physiological: most exposed spp. killed by major changes, e.g. severe rainstorms

--BUT If changes within the range of 150/200-1500 mOsm:

i) Osmoregulation --tidepool fish: gills regulate blood by NaCl transport, reversible!

Or ii) Osmoconforming with Compatible Osmolytes: example of DMSP in algae!

c) Behavioral Avoidance--crevices/under algae

d) Life-cycle: planktonic larvae to recolonize; crucial since major storms can kill off adults.

6. Nutrients for producers: none when tide out; macroalgae cannot absorb from rocks

• Physiology: cyanobacteria & lichens symbiosis: PO₄ from rocks, and

--N2 fixation by cyanobacteria --reason for success in splash/upper zones!

7. Light Intensity: No canopy can develop due to waves

a) Anatomy--symmetrical blades

b) Physiology --UV absorbing pigments: READING on mycosporin amino acids=MAAs

BIOTIC: 8. Food/prey availability--none when tide out , so go dormant

Exceptions: animals with lungs! Know: Littorina; and Surf bird’s adaptation from the video

9. Predator exposure--more intense in LOWER zones

a) Armor-- shells; spines; b) CAMOUFLAGE: e.g., chiton with algae on it

b) Toxins, noxious compounds; e.g., Phaeophytes--vacuoles w/H2SO4, etc.

c) Behavioral avoidance

d) LifeCycle: as before

10. Competition: a) Physiology: anti-competitor compounds

b) Behavior: Territoriality—Anemone clone wars -- acrorhagia battles

ii) Farmer LIMPET patrols territory, removes competitors, slams carnivores!

c) LifeCycle strategies—i) colonize rapidly w/plantonic larvae and grow fast; ii) clone; iii) direct devel.

ECOLOGY: Factors/Adaptations lead to ZONATION: be able to draw and explain zones and KITE diagrams of key factors; typical organisms

--Refuge and harsh-environment hypotheses; how this shows that variable abiotic stresses are harder to adapt to in UPPER ZONES; what rewards accrue to succeeders through COMPETITIVE EXCLUSION!!!

--Intermediate disturbances: role of waves, keystone seastar in enhancing diversity in LOWER zones – prevents Competitive Exclusion

HANDOUT READING--how factors can yield very high productivity; new studies on cooperation

SANDY BEACH COMMUNITY

ZONATION due to similar factors--Upper detrital zone with amphipods, etc.; lower burrower zones; Infauna & Meiofauna

ADAPTATIONS:

1. WAVE Shock: a) Armor: heavy ribbed shells: ribbing to grip sand

b) Behavior: dig deep like geoduck;

c) Physiology--biological clocks e.g. in beachhoppers--how used, why useful

2. Water –dig down to water table or stay under detritus

3. Oxygen--i) lungs -- beach hopper amphipods

ii) Go anaerobic – shallow clams

iii) Stay in water – deep burrowers & migrating “wash riders”

4. Temperature 5. Salinity

} buffered by burrowing to water table

6. Nutrients; 7. Light—diatoms must live low down and near or on surface: clock!!

8. Food availability –bird beaks! VIDEO: how bird beaks are used, how they avoid competition on beach!

9. Predator exposure--Defenses

a) Armor; b) deep burrowing ; c) fast burrowing w/light-weight shell (tradeoff!). Know how foot works!

MUDFLAT COMMUNITY

Zonation—often Wetlands = upper saltmarsh/mangrove, lower open mudflat zones. Why WATER, WAVES not problems!

Factors & Adaptations:

1. Oxygen: vertical profile of oxygen and hydrogen sulfide!

a) Physiology: anaerobic pathways in decomposer bacteria!

(CH2O)x + SO4=---> CO2 + H2S ]and (CH2O)x + NO3----> CO2 + N2 ]

b) Physiology—animals’ hemoglobins

c) Behavior: open burrows with active aeration; H2S toxic (WHY?)

d) mangrove pneumatophores

2. Light, Chemical Energy: Physiol/Biochem adaptations:

--Photo and Chemosynthesis: Sulfur- bacteria!

Photo: 12H2S + 6CO2 + light ---> 6CH2O + 12S + 6H20

2H2S + O2 ---> 2S + 2H20 later: S + 2O2 ---> SO₄

3. Temperature 4. Salinity

} buffered in burrows

5. Food availability: how the unusual Macoma vacuum-cleaner clam outdoes other clams!

6. Predator Exposure:

B. SUBTIDAL Habitats: Temperate Zones

ROCKY ZONES and ZONATION—basic effects of light, waves, currents

a) INSHORE zones; Encrusting or fouling community

b) KELP FOREST in many rocky areas= intermediate zone; holdfast microcommunity: basic factors, food web

c) OFFSHORE Zone: often sediments, some rocks; filterers, scavengers, deposit feeders

KELP FOREST ECOLOGY

How kelp energy enters the food chain; OTTERS as keystone species and history of urchins, orcas

FACTORS/ADAPTATIONS:

1. Waves, Tides, Currents: Holdfast glue; flexible cell walls; etc.

2. LIGHT / Depth: branched, symmetrical blades, phaeophyte pneumatocysts, and PIGMENTS: how red and brown algae compete

3. NUTRIENTS: upwelling, shallow mixing is critical; macroalgae take up nutrients on all surfaces

5. FOOD/PREY—example of sea otter; flexibility of octopod arms

6. PREDATION—DEFENSES: a) Armor: shells etc.•coralline algae

b) Noxious chemicals: •CO—pneumatocyst •Toxins in soft animals •Dynamic camouflage:octopods

c) Behavior: •Behavioral camouflage =decorator crabs

•Escape responses

SUBTIDAL -- SOFT-BOTTOM Habitats

Know basic adaptations/roles of EELGRASS

PELAGIC BIOLOGY

PRODUCERS--know each type, basic features

2) Cyanobacteria: top producers by cell count; much unknown; read on new group = picobiliphytes!

3) Microalgae Protista:

--DIATOMS: general form, SiO2 frustule; radial shape in plankton; #1 top producers by biomass;

--Dinoflagellates: 2 flagella, armor; most photosynthetic but not all

--Coccolithophorids--CaCO3 plates/one cell; READING on functions

--DECOMPOSERS—Eubacteria; Archaea: much unknown

--CONSUMERS—know types, basic features & habitat, what and how they eat:

1) Protista--PROTOZOA

a. Foraminifera--amoeboid, pseudopods through forams in chambered CaCO3 shell; eat bacteria, microalgae

b. Radiolaria--amoeboid, pseudopods, silica skeleton, eat same

2) ANIMALS: know PHYLA as follows, especially CAPITALIZED ones: basic features, how they eat:

• CNIDARIA--carnivores: nematocysts! Medusa and colony gasbags: Scyphozoa, Cubozoa, Hydrozoa

• CTENOPHORA--carnivores--2 sticky tentacles: 8 rows cilia

• MOLLUSCA--head/foot; radula structure, etc. i) Cl. GASTROPODA--heteropods, pteropods

iv) Cl. CEPHALOPODA--radula + beak; carnivores; shelled or reduced or lost shell--e.g., SQUID

• Annelida -- swimming polychaetes
• CHAETOGNATHA--carnivore (jaws) = arrow worms ; eat copepods
• ARTHROPODA subphylum CRUSTACEA; 2 pair antenna at front:

Cl. COPEPODA--filterers with limbs; swimming antennae; #1 herbivores on Earth!

ii) EUPHAUSIDS: KRILL -- shrimplike, but only have swimming limbs

iii) DECAPODS: shrimp

• Echinodermata -- swimming seacucumbers; see later

•CHORDATA subphylum UROCHORDATA--2-siphon filtering system

ii) Cl. THALIACEA--salp form! iii) Larvacea -- tadpole-like forms

• CHORDATA subphylum VERTEBRATA: many feeding types

A. EPIPELAGIC Zone--PLANKTON:

Be able to define these: Phyto- vs. Zoo-; Mero- vs. Holo- plankton

OVERVIEW of PELAGIC ECOSYSTEMS

5 common food chains –know THREE

i) phytoplankton-->herbivore zooplankton ->carnivore (2 or more levels); e.g., diatom-->copepod->chaetognath->fish, etc.

ii) plankton->filter feeder->higher carnivores

iii) MICROBIAL LOOP: up to 25% of energy via producer death/leakage, then bacteria, protozoa-->animals
ADAPTATIONS
1. Density: Sinking bad; need BUOYANCY MECHANISMS

a. Body shape = Increased Friction via high SA/volume ratios--slows sinking greatly, waves suspend like dust motes in air

àHow SMALL SIZE helps; also spines and other thin features !

b. Physiology = Reduce DENSITY--protein, shell problem

(i) Low-density compounds: fat/oil vacuoles in diatoms ; waxes in copepods!!; ii) lighter ions; e.g. squid

(ii) GAS--not so common: Siphonophores (gasbag colonial hydrozoans) as example (Man O'war)

c. Behavior: Motility--all flagellated, ciliated, limbed swimmers

--Scyphozoa (jellies)—weak jet propulsion --riding animals! symbiosis

2. FOOD and FEEDING: --know Copepod & Chaetognath modes

3. PREDATION DEFENSEs:

a) Body cover--shells, spines; gelatinous transparent stealth camouflage! how achieved with microbumps

b) Toxins--red tides: dinoflagellate neurotoxins; animals suffer, or co-evolve resistance!

c) Bioluminescence--startle and slow predators? Burglar alarm!

-- usually Blue or blue-green to penetrate! How made by luciferin/luciferase

d) Behavioral: Escape responses

4. Currents/Eddies & Temperature: plankton swept to wrong thermal habitat: die or

Lifecycle Adaptation: enter dormant stages. How this accounts for BIHEMISPHERE distribution of planktonic species.

5. Salinity --DMSP as key osmolyte: READING on role in climate, possible UV protectant!!

PLANKTON ECOLOGY: know the roles and examples on the global map of the importance of 1) UPWELLING; 2) Shallow MIXING; 3) SPRING/SUMMER; 4) Runoff; 5) Eddies. BLOOM cycles in polar, tropical, temperate seas-- due to light, nutrients, and annual changes

B. EPIPELAGIC--NEKTON

1. MOTILITY: Multipurpose ADAPTATION defines nekton!

Adaptation to staying in light zone; currents; temperature; predation; reproduction

KNOW the two types of turbulence and general solutions:
i) wake turbulence: -- how teardrop shapes help; also golf-ball dimples

ii) flow separation -- how "riblets" help!

*) Anatomical adaptations to Reduce friction & turbulence. EXAMPLES:

i) Sharks-- Skin denticles as riblets; teardrop; also ram ventilation, endothermy, lunate tail

ii) Tunas (Scombrids)—Teardrop; Finlets as riblets?; Smooth mucus; Fin grooves; lunate tail; ram ventilation, endothermy

iii) Humpbacks --reading on tubercles

iv) Dolphins: Teardrop; lunate/truncate tail;; 3-hinged tail with spring energy; endothermy; dynamic riblets

in skin at high speed? BEHAVIOR porpoising & surfing (Bow-wave riding!) & slip-stream riding--how save energy

*) Thrust mechanisms, devices:

i) ROWING/paddling—e.g., duck feet: one power stroke only so inefficient

ii) UNDULATING with CAUDAL FINs: two power strokes—compare tail types

***lunate, truncate, round tails and their features and uses!

iii) JETTING: squid mechanism

iv) FLYING: how wing can give two power strokes using lift /thrust and lift/recovery mechanisms

2. DENSITY/Depth --Buoyancy needed to stay where most food is:

i) CEPHALOPODS: --Low density ions + “pen” instead of a shell in squid

ii) CHONDRICTHYES: cartilaginous bones; Dynamic lift: how this works in a shark with heterocercal fins and snout angle

iii) OSTEICTHYES: swim bladder: how the gas-gland and countercurrent work to trap O2

3. Temperature: a) ECTOTHERMS: Evolution of different temp. optima through protein mutations

Polar: Osmoregulating polar FISH & Antifreeze proteins

b) ENDOTHERMS: lamnid sharks, scombrids, mammals, birds. Use insulation and retes as countercurrent heat-exchangers: how trap heat in fish red muscles, dolphin flippers, etc.

Overheating proble: rete COOLS dolphin testes, can allow heat out through flippers, etc.

*) seals "basking" on land: sunning or escape predation? Reading on re-thinking of this!

4. Oxygen: ram ventilation; blowhole

5. PREDATION-- DEFENSES:

a) Camouflage--Countershading

b) Squid ink--how made, and acts as possible decoy shape!

c) Behavior: Schooling! --Probability factor: how radius of detection favors schooling

-- Escape Maneuvers—flash expansion confuses predator’s limited short-term memory

**How LATERAL LINES of fish work and help coordinate schooling

6. FOOD/PREY--Predation Offenses:

GENERAL PROBLEMS: 1) tiny base of food chain; 2) schooling; 3) hidden prey

CO-Evolution: filter-feeding; corralling/stunning/slashing; and non-visual senses

a) FILTERING: Clupeoids, whale sharks; Mysticeti = baleen whales

i) gill raker mechanism; baleen mechanism & how used

ii) Behavior: Video notes: cooperative humpback bubblenetting

b) CARNIVORY-- scombrids, most sharks, Odontoceti = toothed whales; pinnipeds

i) Antischooling: herding/corralling behavior; mass stunning devices; ii) Non-visual senses

EXAMPLES:

i) SWORDFISH/MARLIN etc: corralling behavior, slashing devices!

ii) THRESHER SHARKS: how tail used

iii) PINNIPEDS: Whiskers as turbulence detectors; reading

iv) SHARKS in general: Electroreception--ampullae on snout. How used, how metal fools

v) ODONTOCETI: Echolocation: produced by sacs/lips, focused by melon, received by jaw fat and inner ear. Versatility in discriminating targets; possible "aural vision" of world

--may also use to stun prey???

vi) SeaBIRDS: Pelican pouch; Albatross Olfactory sensing—DMS smelling and why useful !

REST OF NEKTON MATERIALS will be on FINAL EXAM

READING:

Lecture 14, Feb. 27: --LOVE CODE: what is interesting about mantis shrimp eyes?

Lecture 15, Mar. 2: --SEA URCHIN YIELDS: what is interesting about their spines?

Lecture 16, Mar. 4: --ALGINATES, AGAR, CARAGEENAN: are used how?

--WARMING SEAS DRIVE..why and how are intertidal animals moving north if they can’t crawl far?

Lecture 17, Mar. 6: --NATURE's SUNSCREEN: what is this product based on?

--ENEMY ANEMONE : why do anemone clones often reach a stalemate in the wild?

--FARMER LIMPET--unusual behavior in what ways?

--TOYOTA’s ..car is made of what and why?

Lecture 18, Mar. 9: --WHEN COOPERATION -- what is the 'foundation' species idea here?

--LIFE THRIVES: how do WAVES actually boost productivity of Tatoosh Island intertidal?

--How have DUNE RATS evolved for camouflage?

--AMAZING: what are MEOIFAUNA, recent interest and findings, key examples?

Lecture 19, Mar. 11: --NEW MARINE SPECIES and UPDATE: what new species were found in the Aleutians?

--KELP FORESTS WIDESPREAD: what is this new discovery?

--KILLER in the KELP: what ecological shifts may be happening and why?

Lecture 20, Mar. 13: --OTTER DECLINE affects eagles how?

Lecture 21, Mar. 24; FIELD TRIP

--TRANSPARENT ANIMALS--what is this recent rethinking about salps, etc.?

--KRILL KICK UP--how do they affect ocean mixing?

--FLORIDA RED TIDE: what is the toxin here and an application?

--HOT SPOT -- what was found about the Strait of Juan de Fuca?

LECTURE 22 Mar. 30: --ALGAE ANXIETY; what was found about iron?

--NEW GROUP OF ALGAE: what are picobiliphtes and their significance?

--ROBOT OCTOPUS: what is the bioinspiration here?

LECTURE 23, Apr. 1: --BRISTLING SKIN—what was found about shark hydrodynamics?

--WHEN SUN'S TOO STRONG…what else besides osmolyte role could DMSP be used for? How does it affect climate?

--OCEAN IN A BOTTLE: what is going on with DMSP here?

--DIAGRAMS: adaptations of sharks reduce turbulence

--RED TIDE RISING: can we predict red tides? Why or why not?

LECTURE 24, Apr. 3: --CRABS NOT ONLY—what was found about pain here?

--MOTHER’S SLIPSTREAM helps how?

--WHO ATE ALL—what happened in the current iron-fertilization experiment?

--STRATEGY: FLIPPERS and PRODUCT: what’s the deal with humpback tubercles?

--BIOMECHANICS--SPRING LOADED: how does dolphin tail work?

--SCIENTISTS DISCOVER SECRET… how might dolphin molting help with speed?

--DOLPHIN SKIN--what is the COMPLIANCE aspect about?

--MARINE TECHNOLOGY--is using dolphin biology how?

LECTURE 25, Apr. 6:--GENES HOLD--what is the finding on ANTIFREEZE genes?

--SEALS FORCED…Why do seals sit in the sun?

--HOW DOLPHINS KEEP… their testes cool?

--STUDY REVEALS what about humpback dining?

LECTURE 26, Apr. 8: --BY A WHISKER: how do seals use their whiskers as sensors? How were SealCams used in this study??

--WHALES DRINK SOUNDS: what is the new finding on hearing?

--ARTIFICIAL ARRAYS--how might biomimicry of fish lateral lines be used?

--NARWHAL’S TUSK does what?

--BANG YOU’re DEAD: what is involved here?

--NAVY, WHALE ADVOCATES and OCEANS TOO NOISY: what is the issue with navy sonar testing and other human noise?