LSM3254 Ecology of Aquatic Environments

Learning outcomes:
By the end of this lecture you should be able to discuss various aspects of nekton, including:

The open ocean

What nekton is! Movement and lift Sensory adaptations

Peter Todd Dept of Biological Sciences

Migration Examples of invertebrate and vertebrate nekton

Zones of the marine environment

What lives there?

Artificial/utilitarian classification of marine organisms is generally by habitat and mobility: Plankton: passive drifters Nekton: active swimmers Benthos: bottom-dwellers

Nekton consists of:

Invertebrates:
Cephalopod molluscs Some crustaceans

Vertebrates:
Marine mammals Seabirds Marine reptiles Fishes

General characteristics

Movement
Swimming Active swimming required for: Movement from point to point. Maintaining position against a current (e.g., damsel fish guarding territory in coral reef) Requires well-developed muscular systems High energy requirements Impressive feats of migration (e.g., for feeding; for reproduction)
Movement useful for: Reproduction

Movement

Feeding (finding food and/or remaining in food-rich environment) Protection (predator avoidance) Respiration (aeration of gills) Generate lift

Lift
Nekton generally denser than seawater Need to achieve lift (buoyancy) Static lift Accumulation of materials of low specific density Reduction of heavy tissue Swim bladder D Dynamic lift i Nekton which chase prey actively have generally dispensed with buoyancy aids and maintain lift mainly through constant locomotion e.g. some sharks, tuna

Static lift
Liver

Materials of low specific density Low density lipids (fats or oils) and hydrocarbon (squalene) accumulated in liver and muscle tissues of sharks and some bony fishes
Liver up to 1/4 of the body weight in some sharks Also serve as important stores of energy

Layer of blubber below the skin of whales and seals

Also serve as insulation from the cold

Reduction of heavy tissue Cartilaginous skeleton in chondrichthyians (sharks, rays, skates).

Cartilage lighter than bone

Static lift
Fats, oils and some body fluids only slightly less dense than seawater For many small but active nektonic species expending energy carrying blubber, oil-filled liver, etc. may not be ideal

Swim bladder

SOLUTION: internal gas-filled flotation organ

Swim bladder in bony fishes (osteichthyians) Diverticulum of digestive tract Flexible and extensible containers (cf. cephalopods) Original respiratory function - forerunner of tetrapod lung Maintain neutral buoyancy (static lift) Single or paired bladder:
Physostomatous: connected to pharynx by pneumatic duct (primitive condition); dorsal or ventral connection to pharynx Physoclistous: completely isolated from digestive tract; gas gland and oval window to add and remove gas from bladder.

Static lift
Gas-filled buoyancy organs may be rigid and strong as in some cephalopods - can structurally resist the increased water pressures found at depth e.g. nautilus, cuttlefish il l fi h

Dynamic lift
Heterocercal tail
Sharks (chondrichthyians) lack swim bladder unable to maintain buoyancy and achieve lift by static means alone Some buoyancy from high concentration of low density lipids and hydrocarbon (squalene) especially in liver; and cartilaginous skeleton Compensate with dynamic lift (requires locomotion) Heterocercal tail (dorsal lobe larger than ventral) providing upward propulsion, together with rigid, flattened, laterally-held pectoral fins, acting as hydrofoils, generates lift as fish swims

Cuttlefish has cuttlebone - organ of buoyancy with a number of gas chambers laid down as the animal grows.

The larger upper lobe of a shark's tail cuts the oncoming water slightly before the smaller lower lobe.

Sensory reception
Chemoreception-olfaction e.g., salmon and sharks respond to very low concentrations of odour molecules navigation d i migration; prey d t ti i ti during i ti detection Balance/equilibrium statocysts (invertebrates); otoliths (fishes) able to detect speed as well as gravity - two types of receptor, one detects gravity, other detects acceleration

This creates a wake within a wake, giving the shark both thrust and lift, both forward and upward motion. p

Wilga, C. D. & G. V. Lauder, 2004. Biomechanics: Hydrodynamic function of the shark's tail. Nature, 430: 850.

Vision nekton generally have large well-developed eyes

Some mesopelagic fish have:

Tubular eyes very acute vision in direction that eye is pointed. Retina extends up one side of eye to improve and expand lateral visual field.

Sensory reception
Sound reception Swim bladder of fish:
bristlemouth with tubular eye

Role in detecting vibrations amplifying and transmitting to inner ear vibrations, Sound-production in certain fish - communication, defence, courtship or echolocation

Sound detection in fish:

Inner ear and otoliths (ear stones) Lateral line system detects vibrations in the water

Marine mammals can echolocate and use sound for communication

Odonticeti have a fatty organ at the front of the head called a melon, which focuses emitted sounds from the nasal sacs into a beam ahead of the animal.

Sensory reception
Electro-reception/magneto-reception Some organisms (e.g., sharks) detect weak electric and electromagnetic fields generated by:
Muscle contractions in swimming animals Water currents moving past inanimate objects Earths own magnetic field

The whale produces a beam of f rapidly emitted echolocation clicks to help it navigate and find food.

The clicks bounce back to the j path. whale from objects in its p Sounds are received and conducted through the lower jaw to the middle ear, inner ear, and then to hearing centers in the brain via the auditory nerve.

Sharks, skates and rays have an extensive network of tiny pores or pits on tops of their heads Each pit connects to a flask-shaped jelly-filled ampulla of Lorenzini that is associated with the lateral line system Prey detection Navigation

Tiger shark

Defence and camouflage

Strongest adaptations for defence are those related to achieving fast f f t forward movement in the water column d t i th t l Camouflage nevertheless possible and useful even in open waters of pelagic environment Some species of squid may be transparent, but difficult for most nekton when the body is highly muscular

Ampullae of Lorenzini

Cryptic colouration
Threshold of sensitivity as low as 5 nV/cm. That is 5/1,000,000,000 of a volt measured in a centimeter-long ampulla

Countershading

Countershading
Common in nekton; in many fishes, whales and squid Darker green, grey or blue pigmentation on dorsal surface
Viewed from above, the pigmented upper surfaces blend with the darker background b l b k d below

Migration
Most nekton start life as plankton Spawning sites provide planktonic larvae with access to nursery areas (food and shelter) In certain species, spawning and adult feeding sites may differ, making it necessary to have migrations Migration circuits involves circular movements to:
Spawning grounds Nursery grounds Adult feeding grounds

White or silvery ventral surface

Viewed from below, difficult to distinguish from ambient light coming from the sea surface above

Protection against predators Confuse prey

Generalised migration circuit of migratory nekton

Adult population

Nektonic invertebrates

Spawning site Passive drift as plankton

Nursing area

Giant squid, caught off Spain, Sept.02

Cephalopod molluscs

90kg, 11m long

Squid, cuttlefish, octopus, nautilus comprise a major component of biomass globally Agile swimmers, with complex nervous system Molluscan shell reduced or lost In some regions, vast numbers of small squid (less than 1m in length) form important intermediate links in epipelagic food chains The giant squid (Architeuthis) lives in deep waters
Bathypelagic squid

Funny squid

Antarctic krill (Euphausia superba)

Cephalopod molluscs

Crustaceans
Shrimps/prawns and crabs Some workers regard adult Antarctic krill (Euphausia superba) as nekton Movement:
Prawns and krill use abdominal paired appendages (pleopods) and tail fan (uropods) Certain crabs, use flattened appendages, usually lined with setae (hairs)

Movement:
Water taken into mantle cavity and expelled at high speeds through a nozzel-like siphon (jet propulsion) Siphon can be aimed in any direction for rapid course corrections and for manoeuvring purposes Also use undulating fins on sides of mantle Excellent camouflage.

Swimming crabs (Portunidae)

Light produced by mesopelagic species closely matches background light.

Nektonic vertebrates
Why the clear eye shield?

Marine reptiles

Fishes
Bony and cartilaginous fishes (osteichthyes os-te-ich-they-ez & y y ) chondrichthyes cond-drik-they-ez)

Marine iguanas and saltwater crocodiles

Littoral animals - rarely venture out of sight of land

Sea turtles
Nine species - largest is the leatherback or leathery turtle Long migrations - return to land to lay eggs

Holoepipelagic: spend entire lives in the epipelagic

Most abundant in surface waters of tropics and subtropics

Meroepipelagic: spend part of life cycle in the epipelagic Includes: I l d

Species that live in epipelagic but move inshore or into freshwater to spawn, e.g. salmon, capelin Species that migrate to epipelagic only at certain times to feed, e.g. deepwater fishes

Sea snakes
Laterally flattened tails, which are paddle-shaped Highly venomous (closely related to cobras)
Blue-lipped sea krait

Hard body

Fins - thrust
High speed cruising swimmers (e.g. tuna):
Thin caudal peduncle (minimise drag) Large crescent or forked shape caudal fin (maximise thrust)

Fish swim either by: body and/or caudal fin (BCF) movements - greater thrust and acceleration. median and/or paired fin (MPF) propulsion - generally employed at slow speeds, offering greater maneuverability and better propulsive efficiency.

Fast acceleration and high maneuverability (e g (e.g. grouper):

Thicker peduncle Truncate or rounded tail

Body form - examples of specialists:

Fins Stabilisation of fish

Paired fins (pectoral and pelvic fins) - counter pitch Pectoral fins also used for braking and steering (cf. stiff, inflexible pectoral fins of sharks and tunas used as hydrofoils) Unpaired fins (dorsal and anal fi ) - counter yaw l fins) t and roll

Flat fish (flounder, sole) - bottom specialist

Excellent camouflage g Laterally compressed but look dorso-ventrally flattened One eye migrates to top as fish develops

Deep sea dragonfish (Stomiidae melanostomias)

OCEAN DEPTHS EPIPELAGIC (0-200m) sufficient light for photosynthesis MESOPELAGIC (200-1000m) - dim light insufficient for photosynthesis BATHYPELAGIC (1000-4000m) - no light ABYSSOPELAGIC (4000-6000m) HADOPELAGIC (below 6000m)

Deep water fish

Deep Sea

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Each zone presents different environmental conditions and therefore supports different species communities.
Decreasing light, primary productivity, temperature, food, current strength, environmental fluctuations.

Deep sea fish adaptations

Mostly small-sized (not enough food to grow very big!). Common features of deep sea fishes: - large eyes in the mesopelagic, small below that. - photophores - large mouths with hinged/extendable jaws - sharp teeth - broad diet - small size, lightweight (no scales or spines) -l less muscle, no swim-bladder l i bl dd - soft, weak bones - loss of defensive spines/scales - inactive swimmers.

Declining abundance with depth: 5-10x less organisms at 500m, 10x less at 4000m. - Diversity!??

Increasing nutrients, pressure.

FOOD ACQUISITION
Food capture. Some species have very large mouths and can consume prey larger than themselves e.g. swallowers and gulpers. Stomachs expandable to accommodate prey. No scales, usually black, up 1m long.

Aristostomias sp.

Swallower eel anglerfish anglerfish

Gulper eel

devilfish d ilfi h

swallower gulper bristlemouth

The Malacosteid family (Loosejaws)

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Marine mammals
Marine Mammals fall into 3 orders: Order Si i O d Sirenia - manatees & dugongs. t d Order Carnivora - seals, sea-lions, walruses, otters, bears Order Cetacea - whales, porpoises, dolphins;

They can also be divided another way: Oceanic: whales, seals and sea lions Other marine mammals such as manatees, dugongs (Sirenia) and sea otters (Carnivora) occupy inshore waters at all times Walruses relax and mate on Arctic ice floes. Polar bears spend most of their time on Arctic ice (and may never touch land during their lifetime).

Marine mammals

US FWS

Sirenians (order Sineria)

Monophyletic group with two extant familes: Trichechidae (manatees) 3 species Dugongidae (dugongs) 1 species Wholly aquatic Once heavily hunted US FWS Lips are large and mobile The only herbivorous marine mammal Closely related to elephants (and other sub-ungulates) Now vulnerable to habitat loss and propellers, etc.

Steller s Steller's sea cows discovered in 1741

Georg Wilhelm Steller, physician, and Vitus Bering explorer.

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Or Order or Superfamily

Northern Fur Seal Callorhinus ursinus

Pinnipeds

South African and Australian Fur Seals Arctocephalus pusillus

Harp Seal Phoca groenlandica

(Order Carnivora, Suborder Pinnipedia)

Latin words 'pinna' ('fin', 'flap' o r'wing') and 'pedis' ('foot'),

30,000 killed for fur
Hooded Seal Cystophora cristata

Most found in polar, sub-polar and temperate waters. Share their time between the water and the land (or ice) On land they mate, rest, give birth, and moult.
They are streamlined - nostrils close underwater

60-80,000 killed for fur

Leopard Seal Hydrurga leptonyx

Lots of blubber Underwater, their heart rate slows down and arteries squeeze shut
so that only the sense organs and nervous system get the normal blood flow

Crabeater Seal Lobodon carcinophagus

California, Galpagos and Japanese Sea Lions Zalophus californianus

seals vs. sea lions

Seals Family Phocidae No external ear flaps Cannot rotate rear flippers Swim with rear flippers
NOAA

Sea lions Family Otariidae External ear flaps Rotate hind flippers Swim with front flippers pp

Family Odobenidae
Two subspecies: the North Atlantic and the, larger, the larger North Pacific walrus. Both are usually found above 58 degrees north.

1 species

Odobenus rosmarus

The walrus

NOAA

No external ear flaps but swims with rear flippers p pp

NOAA

Tusks are actually canine teeth (up to 1m long) Males ~ 1500 kg, females ~ 850 kg Benthic feeders - on mollusks (clams) especially

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Sea otters and polar bears (Order Carnivora)

Sea otters (Enhydra lutris)
Smallest of the marine mammals (~45kg) Coastal - max 4 km offshore Often associated with kelp beds (hold on!) Single pup birth in water Rest in single-sex rafts (can be 100s of otters)

polar bear

(Ursus maritimus)
Found in Arctic circle, semi-aquatic Biggest land predator Top of food chain feed on seals p Run at speeds close to 65 km/hr Can swim > 100 km
NOAA

NOAA

Cute!

Also cute!

NOAA

US FWS

Order Cetacea (whales & dolphins)

Order Cetacea (whales & dolphins)

Suborder Mysticeti (baleen whales) 11 species
NOAA

Mysticeti (f M ti ti (from the Greek words M t meaning " th G k d Mystax i "moustache" and K t t h " d Ketos meaning sea monster").

Mostly marine
Includes whales, dolphins, and porpoises Found in all the world's oceans Nostrils are located on the top of the head Specialized forelimbs = flippers Hind limbs and pelvis are very reduced As with pinnipeds, heart rate and blood flow alter when diving. Baleen whales capture prey items by straining water through a series of baleen plates fixed to the upper jaw. These act as filters, collecting food items.

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Suborder Odonticeti (toothed whales) 77 species

Family Platanistidae river dolphins. Only 5 species, e.g. Beiji (or Chinese lake dolphin) now extinct (?) because the Yangtze River has becomes less livable for both prey items and predators.

Dolphin vs. Porpoise

Dolphin Long, sleek body Wave-shaped dorsal fin Rostrum R t Conical teeth Longer life span Porpoise Plump body Triangular dorsal fin Blunt snout Spade-shaped teeth Shorter life span

Chinese lake dolphin

Northern bottlenosed whale

Family Ziphidae (18 species) Beaked whales are deep-water, medium-sized, deep-diving species that are not often seen (species usually known from only a few specimens).

Family Stenidae. 16 species in all including the rough-toothed dolphin, spotted dolphin, long-snouted spinner dolphin, etc.

spotted dolphin bottlenose dolphin

Family Delphinidae (25 species) includes common dolphin, Risso's dolphin, bottlenose dolphin etc. also killer whales and pilot whales.

Family Physeteridae, Genus Physeter: species macrocephalus (sperm whale) and Genus Kogia: (pygmy sperm whale & dwarf sperm whale).

Three main types of feeding behaviour

1. Rorqual whales feed by gulping lots of water containing crustaceans, schooling fi h t h li fishes, small squid etc. Th t ll id t Throat (ventral) grooves (pleats) expand that maximizes the water and food capacity. The jaws are brought together, the throat grooves contract, and the tongue is pressed up, forcing the water to drain out the sides and front of the mouth.

sperm whale What is ambergris?

Left tooth or two twisted upper teeth?

Family Monodontidae Species Monodon monoceros (narwhal) are highly vocal with distinctive horn, up t 3m i l h to 3 in length. th

g beluga

narwhal

Genus Delphinapterous Species Delphinapterous leucas (white whale or beluga).

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2. Gray whales also have throat grooves (but not so many) and filter food through their baleen plates in a similar way to rorquals (except they suck more!). However, first, grey whales dive to the bottom and suck in water, mud, and food (feed (f d on gammarid amphipods, molluscs, crabs and id hi d ll b d polychaete worms).
NOAA

3. Right whales swim at or near the surface of the water and strain th water for small zooplankton such as copepods. t i the t f ll l kt h d Water and food enter the mouth through a gap in the front baleen plates. Water exits through the sides of the mouth and food is caught on the matted baleen fringes inside.

NOAA

NOAA

3 families (3 types of feeding)

Family Balaenopteridae (rorquals) are lunging feeders. This group consists of the Genus Balaenoptera, i.e. minke (pronounced minkee") whale, sei whale ( h l i h l (pronounced " d "say") , B d ' whale ( ") Bryde's h l (pronounced d "broodus") , blue whale, fin whale, plus the Genus Megaptera: humpback whale. The grey whale is on its own in the Family Eschrichtidae, Genus Eschrichtius, Species robustus Family Balaenidae: cruising continuous filter feeders. This group consists of the Genus Balaena: the bowhead whale, and the Genus Eubalaena (right whales): northern right whale, southern right whale, and the pygmy right whale. Grey whale in aquarium!
Plates on right often worn down

Humpbacks feeding (bubble corralling!)

Northern right whale feeding

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Baleen (or whalebone) was used in 19th century corset making and demand helped drive the whaling industry.

Whale-bone traders Probably right whale plates (2-4m)

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