Vertebrates

3 November 2025: Origins of vertebrates 


  • Features of a fish 

    • Paraphyletic

    • A convenient desccirption, not taxonomic ranking

    • Poikilothermic, aquatic chordate

    • Tends to swim, uses gills as main respiratory fins, be bony, be covered in scales, be ectothermic (MANY EXCEPTIONS)

  • Taxnomoic evolution 

    • Major chordate subphyla

      • Urochordata

      • Crainata

        • Chordates with cephalization and bone/cartilage

        • Essentially vertebrates but may lack vertebrate 

        • 8 superclasses

          • Conodonta (extinct)

          • Pteraspidomorphi (extinct)

            • General small (10 – 20cm), but some up to 1.5m

            • Benthic filter feeders

            • Evolved reduced armour, narrower head shield, lateral projections

          • Anaspida (extinct)

            • Small (<15cm), fusiform, compressed

            • Benthic parasitic/detrital feeders: marine → freshwater 

            • Overlapping tuberculate scales

            • Fin-like projections + muscles + internal skeleton → manoeuvrability

          • Thelodonti (extinct)

            • Small (10 – 20cm), fusiform, depressed, large head, horizontal mouth

            • Hypocercal tail, dorsal & anal fins: benthic

            • Forktail form - supra-benthic!

            • Covered with characteristic scales or denticles (like sharks)

            • Stomach

            • Lateral line (first appearance???)

          • Osteostracomorph (extinct)

            • Abundant and diverse

            • Large, anterior, bony shield with eye, nose & pineal openings

            • Ossification of endoskeleton

            • Epicercal tail, body form and paired fins: creates lift

            • Complex brain

          • Myxinomorphi (extant)

            • 70 living species in 1 class: Myxini

            • Temperate / cold temperate oceans, below 30m

            • Restricted to seawater: isosmotic!

            • Sister group to vertebrates: lack even primitive vertebrae

            • No true eyes

            • Mud-burrowing species extremely hypoxia tolerant

            • Predators of benthic inverts and scavengers (burrow into prey)

            • 70 – 200 pairs of slime glands: exude mucous and thread cells

          • Petromyzontomorphi (extant)

            • 38 living species in one class: Petromyzontida

            • Functional eyes, cerebellum, separate ventral & dorsal roots of the spinal nerves (a vertebrate feature)

            • Olfactory and respiratory pathways are separated

            • Freshwater species: may be parasitic

            • Anadromous species: all are parasitic

            • Migrate to upper streams for spawning

          • Gnathostomata (extant)

      • Cephalochordata

  • Overviw of extant vertebrates

    • 55k known living species

      • 27k tetrapods

        • 0.5k marine

      • 28k fishes

        • 16k marine

    • Hagfish versus lampreys

      • Similarities, now thought to be convergent evolution:

        • Eel-like, scale-less, jawless forms

        • Produce pathogen-specific defensive substances: “variable lymphocyte receptors” (vs. antibody proteins in gnathostomes)

        • Tongues posses keratinous, replaceable teeth

        • No stomach

      • Differences

Features 

Hagfish

Lamprey

Mucus production

Yes

No

Fins

Continuous candal

1 or 2 dorsal, caudal

Mouth

Terminal

Sub-terminal

Reproduction

Repeat spawning, direct dev. from egg 

Terminal spawning, larval ammocoetes

Sensory organs

Photoreceptors in head and cloacal region 

Pineal organ, eyes, neuromasts

Gill openings

1-16 

7

Tongue

Biting and tearing

Rasping and sucking

  • Origin of bone in vertebrates

    • Cartilage:

      • Tough, semitransparent, elastic, flexible

      • Glycoprotein strengthened by collagen

    • Mineralized skeleton can be:

      • External (dermal bone) OR internal, derived from cartilage precursors (endochondral)

    • Solid support for attachment of muscles

      • Fast, efficient locomotion → avoid predators and catch prey

    • Storehouse of chemicals (e.g. phosphates) for metabolism

    • Protection


3 November 2025: The arrival of jaws


  • Ancestors of gnathostomes

    • Thelodonts?

      • Stomachs, unlike extant agnathans

      • Scales like teeth and modern shark placoid scales

      • Broad based pectoral fins

    • Osteostracomorphs?

      • Ossified bones around eye

      • Cellular bone

      • Slit shaped gill openings

      • Paired fin structure

      • 2 dorsal fins

      • Epicercal tail 

  • Gnathostomata (extant superclass) (5 classes below)

    • Placodermi (extinct)

      • Bony, ornamented plates over 30–50% of body

      • Big jaws, teeth and large gape: “craniovertebral joint”

      • Quite diverse (200 genera)

      • Dunkleosteus: 1m x 6m or larger!

      • Antiarchs: pectoral fins enclosed in bone-like arms

      • Evolved towards reduced body armour

      • Limitations

        • No replacement dentition

        • Jaw could not generate suction forces

    • Chondrichthyes (extant)

      • 970 living species

      • Defining features:

        • Prismatic calcification of endoskeletal cartilage

        • Pelvic claspers

      • Specialized as marine predators

      • Dominated ancient seas

    • Acanthodii (extinctt)

      • “Spiny sharks”

      • Small and stout

      • Large head and eyes

      • Bony spines before all fins

      • Cartilaginous skeleton

      • Body covered in small scales

      • Water column feeders:

        • Streamlined, round bodies

        • Reduced armour

    • Sarcopterygii (extant)

      • Fleshy, lobed fins

      • Enamelled teeth

      • Cosmoid scales

      • Only 8 extant fishes

      • 26,742 extant species

    • Actinopterygii (extant)

      • 26,891 living species

      • Sister group to Sarcopterygii

      • Monophyletic but no strong derived characters

      • Scales: ganoid, cycloid, ctenoid or absent

      • Spiracle absent

      • Gular plate absent

  • Fish scales

    • 4 main types

      • Placoid

        • Found in Chondrichthyes

        • Dentine covered in vitrodentine (enamel)

        • Richly supplied with blood capillaries

        • Increase in number, not size, as fish grows 

      • Ganoid

        • Found in ancestral Actinopterygii

        • Modified cosmoid scales

        • Dentine covered in ganoine

        • Often rhomboid, articulating peg & socket joint

      • Cosmoid

        • Found in Sarcopterygii

        • Probably a fusion of two placoid scales

        • Cosmine covered in vitrodentine

        • Supplied with blood capillaries

        • Grow from addition of bone underneath

      • Elasmoid

        • Found in Actinopterygii

        • Evolved from ganoid scales: loss of ganoine

        • Fibrous layer (collagen) covered in bony outer

        • Almost completely dermal and grow with fish

        • 2 types: cycloid and ctenoid (with teeth)

    • Ossification of placoderms in skull cases 

    • Chondrichthyans lost the bony skeleton 


6 November 2025: Long in the tooth


  • Class Chondrichthyes

    • 970 living species

    •  Defining features:

      • Prismatic calcification of endoskeletal cartilage

      • Pelvic claspers

      • Specialized as marine predators

      • Dominated ancient seas

    • Subclasses

      • Holocephali (chimeras)

        • 33 spp.

        • Deep water, weird!

        • Gill chamber further forward, no spiracle

        • 1 opercular opening covering 4 gill openings

        • Upper jaw fused to cranium, not protrusable

        • Teeth: 

          • Continuous growth, not replacement

        • Naked skin (modern forms)

        • Claspers on head (also pelvic claspers)

        • Venomous spine in front of 1 st dorsal fin

        • No stomach or ribs

        • Tail often “diphycercal” (pointed) 

      • Elasmobranchii

        • 5 – 7 gill openings

        • Dermal, placoid scales

        • Replacement dentition (derived from placoid scales?)

        • Upper jaw not fused to cranium

        • Ribs

        • Spiracle

        • Several radiations, all extinct except

          •  Division = Neoselachii

            • Overhanging mouth – several jaw modifications

            • Calcified vertebral centra replace unconstricted notochord

            • Basal fin supports: fused, flexible, horny rays support fin

            • Developed sexual dimorphism and internal fertilisation

            • •Electroreception

            • Oil accumulation (buoyancy)

            • 2 subdivisions:

              • Selachii (sharks: 8 orders, 403 spp)

              • Batoidea (rays/skates: 5 orders 534 spp)

              • Differences between the two:

                • Gill slits lateral / ventral?

                • Pectoral fin attached to head?

                • Presence of anal fin?

                • Size and position of spiracle?

      • Subdivision Selachii orders

        • Carcharhiniformes (Requiem sharks)

          • 224 spp.

          • Tropical & subtropical

          • Nearshore

        • Order Lamniformes (Mackerel sharks)

          • 15 spp.

          • Offshore, pelagic

          • Examples

            • Mako

            • White

            • Thresher

            • Basking

        • Order Squaliformes (Dogfish sharks)

          • 97 spp.

          • Successful in North Atlantic, North Pacific & deep sea

      • Subdivison Batoidea orders

        • Rajiformes (skates)

          • 285 spp. 

          • Deep water 

          • High latitude

          • 2 dorsal fins 

          • Long slender claspers 

          • Dorsally depressed 

        • Myliobatiformes (stingrays, eagle rays)

          • 183 spp.

          • Inshore

          • Tropical

          • 0 dorsal fins, tail spine

          • Short, stout, cylindrical claspers

          • Dorsally depressed

    • Body size

      • Relatively large

        • Smallest (17cm): Etmopterus perryi

        • Largest (>12m): Rhincodon typus

    • Habitats

      • Shallow temperate & tropical waters

      • Mainly coastal marine

        • 5% of species in open ocean

        • 5% in freshwater

      • Shallow waters, seldom >3,000m

      • Generally outside environmental extremes

    • Movement and bouyancy in sharks

      • Efficiencies through

        • Integument

        • Fins

        • Swimming type

      • Efficient buoyancy devices

        • No gas bladder

          • …but see Hussain 1989 Indian J. Fish., 36 (3): 266-268!

      • Lightweight skeleton

      • Oil-filled liver

      • Heterocercal tail → creates lift

      • Pectorals to adjust pitch

    • Movement and home ranges

      • Highly mobile, often large home ranges

      • Home range increases with age

      • Different movement patterns:

      • Local

      • Coastal

      • Oceanic

    • Metabolism and growth rates

      • Low metabolic rates

      • Low aerobic scope

      • Low food requirements

      • Variation among species depends on activity

      • Slow growth, long life spans

    • Feeding habits

      • Almost all carnivorous, many apex predators

        • Live prey

        • Scavengers

        • Planktivorous

      • Often nocturnal feeders

      • Strong bite: underslung jaw

      • Teeth for piercing, slicing, crushing

      • Dentition replacement

      • Other key feeding features

        • Tailfin (Thresher sharks: herding & stunning)

        • “Saws” (Sawsharks: slashing).

        • Electricity (Torpedo rays: stunning)

        • Cephalic horns (Manta rays: guiding)

      • Suction in some species

        • From enlarging orobranchial cavity

      • Protrusion of palatoquadrate

        • Improves bites speed and strength

    • Sensory physiology

      • Often nocturnal: rely on non-visual senses

      • High olfactory sensitivity

      • Good vision (esp at night)

      • Mechanoreception

      • Acoustic sensitivity

      • Electroreception

      • Large brain: integration

    • Reproduction

      • Older age at reproduction than actinopterygians

        • Typically mature at 6 – 18 years

      • Internal fertilisation

      • Produce few, successful offspring

        • Low fecundity

        • Large, precocial young

        • Large energetic cost for females

      • Embryonic development & nutrition

        • All nutrition from yolk sac:

          • Yolk sac oviparity

          • Yolk sac viviparity

      • Some additional nutrition from mother:

        • Eating unfertilized eggs or embryos (oophagy or embryophagy)

        • Placenta (placental viviparity)

        • Uterine secretions (uterine viviparity)


7 November 2025: Radiation of ray-finned fishes


  • Class Actinopterygii

    •  26,891 living species

    • Sister group to Sarcopterygii

    • Monophyletic but no strong derived characters

    • Scales: ganoid, cycloid, ctenoid or absent

    • Spiracle absent

    • Gular plate absent

    • 3 subclasses

      • Cladistia (bichirs; 16 spp)

      • Chondrostei (inc. Acipenseridae, sturgeons & paddelfishes; 27 spp)

      • Neopterygii

        • Equal numbers of fin rays as supports in dorsal and anal fins

        • Two types of fin types

        • Orders

          • Lepisosteiformes (Gars): 7 spp.

          • Order Lepisosteiformes (Gars): 7 spp.

          • Division Teleostei (everything else): 

            • 26,840 spp.

            • Mobile premaxilla

            • Neural arches on dorsal side of tail base → uroneural bones

            • Ventral pharnygeal toothplates unpaired

            • Particular arrangement of skull bones

            • Subdivision Osteoglossomorpha (220)

            • Subdivision Elopomorpha (804)

            • Subdivision Otocephala (8344)

              • Superorder Clupeomorpha (364)

              • Superorder Ostariophysi (7980)

            • Subdivision Euteleostei (17422)

              • Superorder Protacanthopterygii (356)

              • Superorder Esociformes (10)

              • Superorder Stenopterygii (391)

              • Superorder Ateleopodomorpha (12)

              • Superorder Cyclosquamata (236)

              • Superorder Scopelomorpha (246)

              • Superorder Lampriomorpha (21)

              • Superorder Polymixiomorpha (10)

              • Superorder Paracanthopterygii (1340)

              • Superorder Acanthopterygii (14,800)

            • Trends in teleostean phylogeny

              • Reduction in bony elements

              • Shifts in position /use of the dorsal fin

              • Shifts in placement and function of paired fins

              • Evolved/advanced caudal fin

              • Gas bladder

                • Arose either as a breathing or buoyancy device

                • Living preteleosts (e.g. gar)  and basal teleosts (e.g. herring) are

                • Physostomous:

                  • Gas released through Pneumatic duct

                • Derived teleosts (e.g. seabass) are Physoclistous: 

                  • Gas exchanged across Rete mirable

                • Loss of dependence on fins for buoyancy → diversification of swimming types

              • A suite of interrelated trends:

                • Increased speed, manoeuvrability and feeding capability

                • No significant loss of defensive structures

                • Diversity of swimming types and morphology

                  • Similar (independent) trends in other groups of fish

            • Skeletal features and scales

Feature

Lower teleosts (e.g. Herring) 

Higher teleosts (e.g. Seabass)

Ossified vertebrae

60-80

20-30

Vertebral accessories 

Many

Few

Bones in skull 

Many

Few

Bones / rays in tail 

Many

Few

Fin rays in paired fins 

Six or more

Six or fewer

Scales

Cycloid (heavy) 

Ctenoid (light)



10 November 2025: Functional morphology + anatomy of fish


  • Senses in fish

    • Sight

      • Eyes: similar to other vertebrates

    • Hearing

      • Good in chondrichthyans and actinopterygians

      • Sound transmits well in water

      • Detection in inner ear

      • Detection based on density differences among tissues

        • Otoliths (in actinopterygians)

          • Hair cells, stereocillia 

        • Gas bladder

      • Sensory epithelium = macula

      • Mechanoreception/neuromasts 

        • Gelatinous cupula displaced by water motion →

        • Cupula moves cilia of hair cells → 

        • Hair cell initiates a change in signals to brain → 

        • Able to filter out background “noise”

      • Two main types of neuromasts

        • Ones that detect slow moving water, located on surface of skin, very easily stimulated

        • Ones that sit inside canals that go down the body, better for fast moving water

          • Lateral line  

    • Sensory input/electroreception 

      • “Ampullae of Lorenzini”

        • Specialized electroreceptors which detect weak electrical fields. 

        • Visible as tiny pores on the skin, enable the animals to find prey buried under sand

        • Navigate using Earth's magnetic field, and follow water currents

        • Two main types

          • Ampullary receptors - receptive to electrical fields produced by biological activity (ex. prey movement) and the environment, can also be used for socail communication and navigation

          • Tuberous receptors - detcted high frequency electric fields, used by fish to detect their own electric discharge 

    • Smell

      • Olfactory chambers

        • Often ventilated by cili

    • Taste

      • Receptors on:

        • Mouth, lips, barbels

          • …also fins and trunk

        • Often clustered into taste buds

  • Maintaining water/salt balance (osmosis let’s goooo)

    • Hagfish

      • Isosmotic, simple kidneys

    • Lamprey

      • Similarities with teleosts (later)

    • Sarcopterygians

      • Similarities with sharks & rays (convergent evolution?)

    • Sharks and rays

      • Slightly hyperosmotic to seawater

      • Excrete salt using rectal gland

      • Concentrate urea and TMAO to maintain high osmolarity

      • Excrete dilute urea

    • Marine teleosts

      • Hypo-osmotic

      • Problem: 

        • Water loss + salt gain

      • Drink water

      • Excrete salt (gills and gut)

      • Excrete scant urine

    • Freshwater teleosts

      • Hyperosmotic

      • Problem: 

        • Water gain + salt loss

      • Avoid drinking

      • Uptake salt (gills)

      • Excrete copious urine

  • Respiration 

    • The buccal pump: chondrichthyes

      • Expand volume of the mouth (buccal) cavity

        • Lower floor of mouth

        • Expand pharynx

      • Sucks water in through the spiracles & mouth

      • Close mouth and force buccal cavity smaller (contract pharynx and raise floor of mouth)

      • Water forced over gills, out through gill slits

        • Water flow is unidirectional and pulsatile

      • Also: RAM VENTILATION

        • A breathing method in sharks where they swim forward with their mouths open, forcing oxygen-rich water over their gills

    • Buccal + opercular pumps: teleosts

      • 2 pumps (give almost continuous flow)

      • At higher speeds: RAM VENTILATION

    • Gills: teleosts

      • Brachiostegal rays/membrane

    • Concurrent exchange

    • Countercurrent exchange

      • Blood along the capillary is always in contact with water that has a higher PO2

  • Circulation 

    • Myxinomorphi

      • Partially open circulatory system

        • Arteries → Sinuses → Veins

      • 4 rudimentary hearts

        • Primary 3-chambered heart

          • 1) Near gills: branchial heart

        • Auxiliary 1-chambered hearts

          • 2) Behind mouth: paired cardinal heart  

            • Re-establish flow

          • 3) Mid-body: portal heart

            • Cardinal vein + intestine → liver

          • 4) End of tail: paired caudal heart

            • Re-establish flow

      • Branchial & portal powered by intrinsic muscle

      • Paired heart powered by extrinsic, skeletal muscle

    • Petromyzontomorphi

      • Partially open circulation, more closed than hagfish

      • Main sinus in branchial region: for blood-gas exchange

      • 1 heart, posterior to gills

      • Cutaneous respiration (CO2 and O2 exchange happens through skin)

    • Chondrichthyes & Actinopterygii

      • Positioned behind gills

      • 4 chambers in series

        • 1. Sinus venosus

          • A reservoir to collect blood

          • Assures easy filling

        • 2. Atrium

        • 3. Ventricle (pump)

        • 4. Conus / bulbus arteriosus

          • Conus, muscular in sharks

          • Bulbus, elastic in bony fish

      • Sinoatrial and atrioventricular valves maintain unidirectional flow


10 November 2025: Tetrapod invasion of land


  • Challenges of moving onto land

    • Features central to the evolution of tetrapods

      • Pectoral (and pelvic) fins

      • Respiratory system

      • Circulatory system

      • Also: Reproduction, digestive system, sensory system

    • Main view

    • Alternative views

      • Movement not limiting

        • Many extant teleosts can move over land

        • Crutching (like mudskippers) may have been the common method

      • Respiration not limiting

        • Some advantages of air breathing

        • Air was higher in O2 at the time

      • Terrestrial reproduction can be a strategy to avoid predation

        • A force driving fish out of water?

      • Feeding in air requires much morphological modification

        • Fish remained tied to water by need to feed?

  • Class Sarcopterygii

    • Key traits

      • Enamelled teeth

      • Fleshy, lobed fins

      • Cosmoid scales

      • 26,742 extant species

        • Only 8 extant fishes

    • Subclass: Coelacanthimorpha

      • Appeared in the Devonian, max diversity in the triassic

      • 2 living species (considered extinct until 1938)

      • 3-lobed tail supported by hollow spine

      • Unconstricted, unossified notochord

      • Double gular plate

      • Spiny dorsal fin

      • Cranio-vertebral joint!

    • Subclass: Dipnotetrapodomorpha

      • Dipnomorpha (includes lungfish: 6 living species, all freshwater)

        • All living spp. are FW: 1 Australian, 1 S. American, 4 African

        • Once considered origin of tetrapods

      • Tetrapodomorpha (all extinct except infraclass tetrapoda)

        • Large predatory fishes

        • Symmetrical tails (functioning swim bladder?)

        • Peaked in late Devonian, extinct in Permian...except tetrapods

        • Robust limb skeleton: hip / shoulder girdles and rotational shoulders

        • Could be large: up to 6m in Rhizodontiformes!

        • Dorsally placed eyes

        • 1 pair of external nostrils (incurrent)

          • Choana (excurrent nostrils moved to palate)

        • Many thought to be ambush predators 

  • Water versus air as a respiratory medium

    • Breathing frequency

      • 1 per 3 -10 mins in obligate air breathers

      • 1 per HOUR for water breathers with lungs

      • Fewer breaths = fewer trips to the surface

    • Water

      • High density (heavy) 

      • High viscosity 

      • Low O2 areas

      • Available everywhere 

      • Easy to expel metabolic wastes 

    • Air

      • Low density (1/800th water)

      • Low viscosity (1/30th water)

      • O2 always available

      • Requires surface access

      • Hard to expel metabolic wastes

    • Bimodal breathing  the best of both worlds

    • Air is more O2 rich: 21% O2 in air vs <1% in water

    • Diffusion from air to blood is also faster

  • Air breathing fish

    •  > 370 known spp (49 families): capacity to obtain O 2 from air

    • Has evolved many (~50?) times

    • Aquatic vs. amphibious

    • Facultative vs. obligate

    • Air-breathing organs:

      • Derived from gut (lungs, gas bladder, stomach, intestine)

      • Head & pharynx (gills, mouth, pharynx, opercles)

      • Skin

    • In higher actinopterygians, lung → gas bladder

  • Origins of lungs

    • Air breathing

      • To cope with seasonal dryness?

      • Exploit new habitats / release competition?

      • To survive low O2 waters?

        • Lungs probably evolved in marine species- not usually hypoxic

        • Habitats of lungfish not necessarily hypoxic

      • Farmer (1999): To avoid myocardial hypoxia?

        • Exercise stimulates air breathing more than aquatic hypoxia

        • Death from exercise results from heart failure

        • O2 sensors afferent to gill

        • Early fish active, high O2 environment

    • Secondary loss of lung?

      • Air access not possible at depth

      • Aerial predation

  • Origins of separate circulation

    • Separation of circulation

      • To survive low O2 waters?

      • Atrium fully divided

      • Ventricle functionally divided

      • Oxygenated blood → reduced gills

      • Systemic blood → gills then lung


13 November 2025: Return to the ocean I


  • Amphibian respiratory system

    • Amphibians: Tetrapods with aquatic eggs

    • Permeable skin and gills

    • Lungs (?)

    • Simple sacs divided by ridges

    • May simply supplement cutaneous respiration

    • Positive pressure breathing: buccal pump + elastic recoil

  • Marine reptiles 

    • Successful in the mesozoic

      • Sauropterygians (plesiosaurs)

      • Ichthyopterygians

      • Mosasaurs

      • Sea Turtles

    • <70 extant spp

    • Tropical / subtropical

    • Three orders

      • Squamata (snakes/iguanas)

        • Snakes

          • True sea snakes (~50 spp.): fully marine

          • Ovoviviparity

          • Sea kraits (~5 spp.): some terrestrial needs

            • Digestion on land

            • Ovoparity

          • Highly venomous

          • Coastal tropical, Indian & Pacific Oceans

          • Excellent swimmers and divers

        • Iguana

          • 1 sp. (Galapagos)

      • Testudines (sea turtles)

        • 7 or 8* spp:

          • Green

          • Flatback

          • Loggerhead

          • Hawksbill

          • Olive Ridley

          • Kemp’s Ridley

          • Leatherback

          • * Some suggest black sea turtle as an 8th species

        • Distinguish spp. by head & shell

        • Circumglobal, tropical

        • Mainly coastal (except leatherback)

        • Good swimmers: foreleg paddles

        • Come ashore to lay eggs

      • Crocodylia (crocs)

        • 2 species:

          • Crocodylus acutus (American Crocodile)

          • Crocodylus porosus (Saltwater Crocodile) 

    • Reptile respiratory system

      • Skin is nearly impermeable to O2

        • But… cutaneous respiration in seasnakes

      • Increased reliance on lung as respiratory surface

      • Lung volume constant

      • Subdivision increases

      • Negative pressure breathing (aspiration pump):

        • Uncouples feeding and breathing

        • Requires thoracic cavity

      • TIDAL VENTILATION

      • Left to right shunt

    • Adaptations

      • Usually v. good swimmers

      • Good divers (esp. turtles)

      • Cutaneous respiration (seasnakes)

        • Do not become anaerobic despite up to 2h long dives

        • Lung extends full length of body

        • Posterior portion oxygen store

        • Small, thin scales and flattened body

      • Cuticle impermeable, but…

        • Salty foods

        • Accidental ingestion of seawater

 Kidney cannot produce urine more concentrated than seawater

  • Salt glands- to excrete excess salt

    • In marine lizards:

      • On head

      • Empties into nasal cavity

      • Ridge prevents re-swallowing

      • Sudden exhalation to expel (sneeze)

    • In sea snake:

      • Base of the tongue

      • Empties into the oral cavity

    • In turtles:

      • In orbit of eye

      • Empties into posterior corner of orbit

    • In Crocodiles:

      • Distributed over the surface of the tongue


  • Sea birds

    • 5 orders

      • Sphenisciformes

        • 17 spp.

        • Penguins

        • All seabirds

        • Southern hemisphere*

          • *Galapagos penguin just north of the equator

        • Flightless

        • Feet → rudder

        • Wings → fins

      • Procellariiformes

        • 125 spp.

        • Albatrosses, petrels, storm-petrels, fulmars and shearwaters

        • All seabirds

        • Tubular nostrils

        • Good sense of smell

      • Pelecaniformes

        • 65 spp.

        • Pelicans, frigatebirds, gannets, boobies, cormorants, anhingas

        • All waterbirds

        • All seabirds, except anhingas, some pelicans and some cormorants

        • All four toes are webbed

        • Salt gland enclosed within orbit

        • Nostrils are slit like, nearly closed or absent

      • Charadriiformes

        • 128 spp. + ~200 shorebirds

        • Skuas, jaegers, gulls, terns, auks, guillemots, puffins, shorebirds & skimmers

        • Mostly seabirds except shorebirds

      • Ciconiiformes

        • Herons, egrets, storks, ibis, spoonbills

        • May feed along the shoreline, but not ‘seabirds’

    • Bird respiratory system

      • Flight + endothermy

      • High metabolic demands

      • Insulation

      • HIGH O2 DEMAND!

      • Tidal ventilation of air sacs → Unidirectional ventilation of “PARABONCHUS” (lung)

      • Cross-current blood flow

    • Adaptations to life at sea

      • Energy management

        • Weight reducing adaptations

        • High MR and endothermy

        • Specialised lungs

      • Salt management

        • Feed in salt water

        • May avoid saltwater ingestion

        • Nasal salt glands- above eye

        • Connects to nasal cavity

        • Preening gland / waterproofing

      • Locomotion and feeding

        • Wings for:

          • Underwater swimming

            • E.g. penguins, cormorants

          • Flying vast distances

            • E.g. albatrosses

          • Flying fast and agile, close to shore

            • E.g. auks and puffins

        • Bodies can be streamlined for swimming underwater

        • Webbed feet

        • Bills adapted to prey type and feeding mechanism

        • Colouration

          • Can be cryptic… or not

      • Life history

        • Form large colonies

        • Large, long life, deferred maturity, small clutch size, extended chick period… due to energy limitation?


13 November 2025: Return to the ocean II


  • Marine mammals evolution and taxonomy 

    • Order Sirenia

      • Includes manatees & sea cows

      • 4 sp.

      • Arose in the Eocene (50 mya) in the Sea of Tethys

      • Evolved from the Proboscideans (elephants)

      • Main features

        • Herbivores

        • Very low metabolic ratergrgthth

        • Tropical / warm shallow water

        • Heavy bones

        • Flipper like forelimbs

        • Reduced hindlimbs

        • Blubber and sparse hair

        • Fleshy, almost prehensile lips

        • 2 families:

          • Trichechidae (manatees)

            • 3 sp.

            • New World

          • Dugongidae (dugongs)

            • 1 sp. (+ Steller’s Sea Cow (extinct))

            • Indo-Pacific

    • Order Carnivora

      • Includes seals, sea lions & walruses

      • Pinnipedia 33 sp.

        • From late Oligocene (27 – 25 MYA)

        • 3 monophyletic families:

          • Otariidae (eared/fur seals & sea lions)

            • 14 sp.

            • ​​Ability to rotate pelvis (walking)

            • Small external ear flaps (pinnae)

            • Dense fur for insulation

            • Long coarse guard hairs

            • Thick under-fur to trap air

            • Large fore flippers for propulsion

            • Sexual dimorphism

          • Phocidae (true or earless seals)

            • 18 sp.

            • Unable to rotate pelvis: move by undulating body

            • No external ear flaps

            • Blubber: insulation

            • Pelvic flippers: propulsion

            • Pectoral flippers: stability & steering

            • Excellent divers

          • Odobenidae (walruses, 1 sp.)

            • 1 sp.

            • Previously diverse, two subspecies today

              • Otarioidea (O)

              • Phocomorpha (P)

            •  Ability to rotate pelvis (O)

            • No external ears (P)

            • Blubber for insulation (virtually hairless) (P)

            • Pelvic / pectoral flippers for propulsion (O & P)

            • Large canine tusks (Neither)

        • Semi aquatic

        • Carnivores

        • Fur/ hair and blubber

        • Very sensitive whiskers (vibrissae)

        • Mobile neck

        • Swim side to side like fish (lateral undulations)

        • Ancestral origins of pinnpieds 

          • Enaliarctos – the earliest known pinnipeds

            • Mid-Oligocene – late Miocene (~30 MYA) from California and Oregon

            • Aquatic features

              • Distinct, highly modified flippers

              • Streamlined

              • Reduced tail

            • Still retained many terrestrial features

              • Inner ears adapted for hearing in air

              • Heterodont dentition: must bring prey to shore to handle + chew it

            • A coastal species

            • Similar to sea otter

            • Ancestral to all pinnipeds?

            • Only in Pacific

          • Puijila darwini: Rybczynski et al. (2009)

            • Late Oligocene  early Miocene, Canadian lakes

            • Similar to land dwelling arctoids

              • No flippers

              • Long tail

              • Proportionally long limbs like a skunk or marten

            • A swimmer

              • Its all in the muscle attachment sites

              • Long shoulder blades (extend down back)

              • Big shoulder muscles

              • Large muscle (teres) that allows rotation of shoulder

              • Robust forelimbs

              • Enlarged webbed (?) feet (flattened finger ends)

            • Not an otter

              • Hands & feet proportions are wrong: big hands unlike otters

              • Long first finger, long phalanges

              • Shorter, slimmer tail not used in swimming

              • Uses all four limbs to swim (otters use only back and tail)

          • Unspecialized! Arctic! Freshwater!

      • Marine otters 2 sp.

      • Polar bear 1 sp.

    • Superorder Certartiodactyla

      • Includes whales, dolphins & porpoises

      • Cetacea (Order) 90 sp.

        • Arose in Eocene (53- 54MYA) in Sea of Tethys

        • Entire life in water

        • Blubber, almost hairless

        • Shortened neck with fused vertebrae, no hind limbs, powerful tail (horizontal flukes), paddle-like forelimbs

        • Blow holes

        • Specialised ear bones and semicircular canals

        • 2 major groups: 

          • Mysticetes

            • Feeding

              • Mouth is flexible, capacious

              • Lack teeth: have baleen plates

              • Eat small fish or krill

            • 2 blowholes

            • Small eyes point sideways

            • Major families:

              • Family Balaenopteridae: fin, humpback

              • Family Balaenidae: right, bowhead

              • Family Neobalaenidae: pigmy right

              • Family Eschrichtiidae: gray

          • Odontocetes

            • “Toothed whales”

            • Peg like, uniform teeth

            • Eat fish and mammals (fast)

            • High frequency sound:

            • Echolocation

            • Communication

            • Melon on skull

            • Fused nostrils → one blowhole

        • Evolutionary origins

          • Artiodactyls (even toed ungulates):

            • Deer

            • Antelope

            • Camels

            • Pigs

            • Giraffes

            • Hippos 

    • Extinct groups

      • Desmostylia (related to elephants), Kolponomos (marine bear- like carnivoran), Thalassocnus natans (aquatic sloth) 

  • Respiration in marine mammals

    • Similar to birds

    • Completely divided

    • No cutaneous respiration

    • Mammalian lungs

      • Back to tidal lungs again

      • Dead space = 1/3 – 1/20th tidal vol

      • 5% body volume

      • Finely divided → V. high SA

      • Surfactants prevent bubbles

      • Negative pressure ventilation

  • Marine mammal adaptions 

    • Diving

      • Record dive: 137.5 min, 2992m (Cuviers Beaked Whale)

      • Challenges: O2 decrease, lactate build-up, pressure change, changes in gas chemistry

      • Heart:

        • Similar size & structure to terrestrial

        • Glycogen stores

        • Aortic bulb

        • Retia mirabilia

        • Myoglobin-rich muscles

        • Increased haematocrit

      • Blowhole muscles

      • Respiratory system:

        • Collapsible chest

        • Reduced lobulation

        • Supported trachaea

        • Blood buffering

      • Diving response:

        • Decline in heart rate

        • Regional vasoconstriction

        • Reduction in core temperature

        • Regional reduction in metabolic rate

    • Senses

      • Sound travels quickly underwater, light readily attenuated

        • Sound used for communication & echolocation

      • Low freq: long distance

      • High freq: fine resolution

      • Sound production

        • Larynx (pinnipeds)

        • Inflatable throat pouches (pinnipeds)

        • Phonic lips + melon (cetaceans)

      • Sound reception

        • Auditory canal (pinnipeds)

        • Lower jaw and throat (cetaceans)

    • Salt management

      • Have lungs, but not immune to problems of high salt:

        • May ingest water accidentally

        • Diet is salty (esp. invertebrates and plants)

    • Internal fluids are hypoosmotic to seawater

    • Derive water from food-preformed and metabolic

    • Large, reniculate kidneys-excrete concentrated urine