evolution of invertebrates and vertebrates Flashcards
what is an invertebrate
- lacks a spinal chord and backbone
- lacks bones
6 evolutionary events led to the evolution of metazoa
- multicellularity
- extracellular digestion
- nervous system
- middle germ cell layer
- bilateral symmetry
- through-gut
evolution of multicellularity
3 theories
- symbiotic theory, different protozoa join together as symbionts
- colonial theory, asexual reproduction of cells that remain together
- cellularisation, multinucleate protist evolves cell membranes around its nuclei
evidence for colonial theory, evolution of mulitcellularity
- choaloflagellate is a protists very similar to collar cells
- proterospongia is accumulation of choaloflagellate cells
- sponge contains collar cells
- none of these are closely related to each other
embryonic development
- embryology recapitulates phylogeny
- blastea = hollow ball of cells
- cells differentiate and rearrange
- blastula forms
- creation of multiple cell layers through invagination or ingression
- gastrula = multilayered embryo
- each cell layer becomes different cell types
diploblastic organisms
- ectoderm = outer layer, differentiates into epithelium etc
- endoderm = inner layer, differentiates into gut lining etc
triploblastic organisms
- ectoderm = outer layer, differentiates into epithelium etc
- endoderm = inner layer, differentiates into gut lining etc
- mesoderm = middle layer, forms muscles, organs etc
non-bilateria properties
- asymmetrical invertebrates
- include basal groups like sponges
- primarily marine
- covered in microscopic pores
- cellular level of organisation, lack tissues and organs
- diploblastic
- sessile
- defensive chemicals
structure of sponges
- pinacoderm = outer layer of cells, pinacocytes = wide and flattened
- choanocytes = collar cells, line central atrium
- gelatinous non-living matrix in central atrium
- totipotent archaeocytes/ameboid cells within matrix, involved in feeding
- spicules
- porocyte = pore cell
sponges, spicules
- small SiO2 or CaCO3 structures
- skeleton-like structural support
- secreted by specialised sclerocytes which fuse, lay down spicules then pull apart
- megascleres support sponge
- microscleres support sporocytes
sponges, digestion
- intracellular
- flagella of choanocytes beat, creating a current to draw in water through pore cell for filter feeding
- food particles trapped in mucous around microvili that make up collar of choanocytes
- water exits through osculum (larger pore)
sponges, reproduction
- external fertilisation
- choanocytes can differentiate into gametes in breeding season, eggs often retained and larvae released
- planktonic larval form drift around in water column
4 phyla in non-bilateria group
- poripheria (sponges)
- placozoa
- Cnidaria
- Ctenophora (comb jellies)
placozoa
- arose from the evolution of extracellular digestion and true epithelium, advantageous for growth and predation
evolution of the nervous system
- gave rise to neuralia clade
- gave rise to cnidaria
- organisms can swim and respond to the environment
Cnidaria structure
- epidermis contains epithelialmuscular cells, totipotent interstitial cells, cnidocytes (stinging cells), mucous gland cells, sensory cells that make up nerve net
- mesoglea = gelatinous matrix in between
- gastrodermis lines gastrovascular cavity and contains enzymatic gland cells, nutritive muscular cells, mucous gland cells and some nerve cells
- 2 morphs
cnidaria, 2 morphs
- polyp, sessile, tentacles pointing upwards
- medusoid, tentacles pointing downwards
- species often travel through both morphs in their life cycle
phyla Cnidaria, class hydrozoa
- marine and freshwater
- colonial organisms
- polymorphic polyps
- cnidae only present on epidermis
- alternation of generations
Cnidaria properties
- anemones, corals, hydroids, jellyfish etc
- once thought to be radially symmetrical but now thought to be bilaterally symmetrical (cilliated groove down middle of animal on inside), so possibly not in non-bilateris
- all have stinging cells, cnidocytes
- mostly marine
- tissue level of organisation
phyla cnidaria, class scyphozoa (true jellyfish)
- all marine
- medusoid stage dominant in life cycle
- thick mesoglea
scyphozoa (true jellyfish) life cycle
- adult medusa is dioecious (either male or female)
- external fertilisation forms ciliated planula larvae that swim around
- planula larva settle and develop into scyphistoma
- scyphistoma undergoes strobilation releasing young jellyfish (ephyrae)
phyla cnidaria, subclass octocorallia
- soft octocorals
- 8 pinnate (feather like) tentacles
- 8 longitudinal septa
- thick mesoglea
- internal calcium skeleton
phyla cnidaria, subclass hexacorallia
- 6 tentacles
- stony sclerotinia corals
- secrete calcium skeleton that the coral sits on top of
- have spriocysts (modified cnidocytes used to catch prey)
Ctenophora phyla
- comb jellies
- younger lineage than placozoa and cnidaria but less complex, possibly shows secondary simplification
characteristics of eubilateria
- bilateral symmetry
- triploblastic
- organs
- centralised nervous system with ‘brain’
- cephalisation
coelomate phyla
- vast majority of eubilateria
- Spiralia, arthropoda, chordata etc
- coelom = fluid filled cavity bounded on both sides with embryonic mesoderm
acoelomate phyla
- lack coelomic cavity
- e.g. Platyhelminthes
Platyhelminthes
- basal phylum of eubilateria
- acoelomate
- varied group, no defining synapomorphies
- incomplete gut, one hole (secondary simplification)
- advanced osmoregulatory organs, protonephridia
pseudocoelomate phyla
- lack true coelomic cavity, only bounded by embryonic mesoderm on one side
- e.g. Rotifera, Nematoda, Priapula
Protostomes
- determinate development, cell fate is fixed from embryo
- presence of a coelom formed through schizocoely in development
- includes spiralia, ecdysozoa etc
Spiralia
- protostomes that undergo spiral cleavage, cell twists when it undergoes cell division
- not a true ranked taxon
- includes Mollusca, annelida, platyhelminates etc
taxon ecdysozoa
- has an exoskeleton that is often calcified
- moults cuticle at least once in life cycle to be able to grow (ecdysis)
- e.g. Phylum arthropoda
Phylum arthropoda
- non-calcified exoskeleton secreted by epidermis
- procuticle made of chitin
- epicutlicle, waxy lipoprotein kayer for protection
- segmented body with a pair of jointed legs at each segment (tagmatisation)
- complex gut with specialisation
- compound eyes
subphyla crustacea
- arthropods
- e.g. crabs, lobsters
- head and thorax fused (cephalathorax), covered with calcified carapcace
- 2 pairs of antennae
- nauplii larvae
subphyla myriapods
- arthropods
- e.g. millipedes, centipedes
- cannot close spiracles so exclusively found in damp environments
subphyla hexapoda
- arthropods
- e.g. class insecta
properties of class insectica
- wings (if present) on last 2 segments of thorax
- thorax segments prothorax, mesothorax, metathorax
- tympanal organs
- tracheal system for gas exchange
subphyla chelicerata
- arthropods
- include arachnids
lophotrochozoa taxa
- 5 phyla that produce a larvae type called tropophore and possess a lopophore
- platyhelminthes, rotifera, nemertea, annelida, mollusca
phylum annelida
- includes earthworms, leeches, ragworms, sipubunculans
- most long and thin, adaptation for burrowing
- specialised through gut with regionalisation
- closed circulatory system with blood vessels (as long and thin)
- defining feature is bristles, chaete
- majority have metameric segmentation
annelida, metameric segmentation
- body comprised of many repeating units
- protostomium = first segment, contains sensory structures
- peristomium = second, has mouth
- metameric segments
- pygmidium = fourth segment, contains anus, organism grows from here
properties of phylum mollusca
- includes snails, slugs, cephalopods, bivalves
- very diverse
- open circulatory system, haelocoel
- visceral mass, organs concentrated into hump
- mantle = sheet of tissue covering visceral mass
- metanephridia = complex kidney-like filtering organs
- e.g. class gastropoda
class gastropoda, shell
- vast majority have coiled (typically clockwise) shell containing visceral mass
- useful for predator evasion
- size constrain means they have lost duplicate organs
- attached to shell via collumellar muscle, enables it to contract into shell
- torsion, internal organs rotated 180 degrees, anus is next to head
- some organisms such as nudibranchs have no shell, so have detorted
class gastropoda
- molluscs
3 recognised groups - prosobranchs (marine)
- opisthobranchs (marine)
- pulmonates (terrestrial)
gastropoda, radula
- radula = toothed tongue used for feeding
- secreted from radula sac
- rasping mechanism
- supported by muscular odontophore
-can be modified into harpoon drill etc - strongest biological material
Deuterostomes
- undergo radial cleavage (cell does not twist during cell division)
- development is indeterminate/regulative
- coelom formed through enterocoely (pouches form coelom)
- 1st pore formed anus, 2nd pore formed mouth
- includes Echinodermata, hemichordata and chordata
Phylum Echinodermata
- include starfish, brittle stars, sea urchins etc
- majority marine
- no head or circulatory system (secondary simplification)
- photoreceptors
- adults have pentaradial symmetry, larvae are bilaterally symmetrical
- water vascular cavity with fluid-filled cavity derived from coelom
Echinodermata anatomy
- dorsal anus, ventral mouth
- 2 stomachs, cardiac and pyloric
- pyloric stomach branches to the pyloric caeca, with papulae to increase surface area
- madreporite = opening that filters water into vascular system
- pedicellariae = extension of water vascular system to clean surface of animal
Phylum hemichordata
- halfway between echinodermata and chordata
- lack a notochord, but have a dorsal hollow nerve chord
- bilaterally symmetrical
- pharyngeal gill slits
- open circulatory system
- complete complex gut
properties of phylum chordata
- non-vertebrates and vertebrates, all non-vertebrates are marine
- notochord = dorsal flexible rod of tissue derived from embryonic mesoderm
- dorsal hollow nerve chord
- bilaterally symmetrical
- pharyngeal gill slits
- postanal tail
- complex gut
- Urochordata, cephalochordata and craniates
paedomorphosis
- larvae become sexually mature before reaching adult form
- neoteny = retention of larval/embryonic characteristics past reproductive maturity
- progenesis = accelerated development of reproductive organs relative to somatic tissue
- theory of how chordates evolved
subphyla urochordata/tunicates
- colonial, solitary or pelagic
- e.g. class Ascidacea
class Ascidiacea
- urochordate
- sea squirts, sessile filter feeders
- heart
- large ciliated pharynx with gill slits (stigmata)
-endostyle (thyroid-like) secretes iodine rich mucous net that traps and digests particles - produces pelagic ‘tadpole’ larvae with notochord present in tail
- notochord degenerates when larvae undergoes metamorphosis
cephalochordates
- chordates that have all chordate features present in adults
- e.g. branchiostoma
properties of craniates
- chordates that have a cranium (skull)
- neural crest
- raised metabolism
- heart with at least 2 chambers
- haemoglobin in red blood cells
- kidneys
neural crest
- embryonic source of many unique craniate characteristics
- pluripotent
- forms peripheral nervous system and myelin sheath
- migrates out of neural plate to form autonomic nervous system, skull, bones etc
- neural plate fuses into neural tube, forms central nervous system
fossil origins of vertebrates, conodonts
- ‘cone teeth’
- abundant over 300mya
- no jaw
- soft, slender bodies
- probably hunters
craniates, class myxini
- hagfish
- least derived surviving craniate lineage
- cartilage skull
- lacks a jaw
- lack vertebrae
- snake like swimming through muscles pushing against notochord
- small brain, eyes and ears
- nasal opening connects with pharynx
- keratinous tooth like formations
- mostly bottom dwelling scavengers
- water-absorbing slime glands, slime repulses/suffocates
properties of vertebrates, vertebrae
- craniates that have a vertebral column
- some vertebrates have vertebrae made of cartilage
majority have vertebrae that enclose the spinal chord and take up role of notochord - supports body
- protects nervous system and brain
- can grow and repair
origins of bones and teeth
- vertebrate skeleton originally evolved as a structure made out of unmineralised cartilage
- mineralisation may have begun in mouth as feeding mechanisms transitioned from filter feeding to predation, as they then needed a way to break down particles
- earliest mineralised structures found are conodont teeth
- armour is then derived from dental mineralisation
- the endoskeleton then becomes mineralised, starting from the skull
fossil origins of craniates, haikouella
- emerged 530mya in the cambrian explosion
- not a true craniate as no skull or ear organs
- suspension feeders
- large well formed brain
- small eyes
- muscle segments along body
- respiratory gills in pharynx
fossil organs of craniates, Myllokunmingia
- more advanced chordate
- regarded at true craniate as has ear and eye capsules as part of skull
other properties of vertebrates
- skull
- well developed circulatory system
- internal organs suspended in coelom
- 3 part brain (forebrain, midbrain, hindbrain)
- duplication of hox gene complex
- neural crest cells
- underwent another gene duplication after branching off from craniates, transcription factor (DIx genes), possibly lead to innovations in nervous system and skeleton
- aquatic vertebrates have fins stiffened by fin rays
class petromyzodontia
- lampreys
- oldest living vertebrate lineage
- majority freshwater
- jawless (agnatha)
- development of an oral hood with keratinous teeth
- cartilage skeleton, no collagen, stiff protein matrix instead
- notochord persists in adults
- cartilagenous pipe around notochord, partially encloses nerve chord
- pairs of cartilage projections related to vertebrae extend dorsally
petromyzodontia feeding mechanisms
- larvae are freshwater suspension feeders
- most adults are parasitic on fish
- round jawless mouth with rasping tongue digests blood and tissues from rasping wound on fish
- anti-coagulant secreted from oral gland
later fossil vertebrates
- emerged during Ordovician, Silurian and Devonian
- paired fins
- inner ear, 2 semicircular canals for balance
- no jaw but muscular pharynx to suck in other organisms/detritus
- armoured, mineralised bone and spines
Gnathostomes, properties
- vertebrates that have jaws
- diverse, outnumber agnatha
- additional replication of hox genes (4 clusters)
- other gene clusters duplicate, increasing genetic and developmental complexity
- enlarged forebrain, complex olfaction and vision
- aquatic gnathostomes have a lateral line system ( sensitive to vibration, early versions in some jawless vertebrates)
advantages of jaws
- active predation, can capture and bite prey
- herbivory
- gripping and slicing food, especially when paired with teeth
- chewing facilitates breakdown and digestion of food
- mouth brooding
- nest building
- mate grasping
- improved gill ventilation
evolution of jaws
- from branchial arches, skeletal rods that support gills
- first gill arch evolves into upper and lower jaw
- second gill arch supports/suspends jaw
- evolution of hyoid bone that supports tongue
gnathostomes, fossil history
- emerged mid-Ordovician ~470mya
- paired fins and tail, jaws with large bite force
- placoderms = earliest gnathostomes, plated/armoured, became extinct in early carboniferous E.g.
Dunkleosteus - acanthodians = marine and freshwater organisms in Devonian era, probably sister group to the bony fish (dermal operculum, hyoid attachment, true dermal teeth)
gnathostomes, Chondrichythyes class
- cartilage skeletons (secondarily derived, traces of bone found)
- skeletons often impregnated with calcium
- sharks , rays and skates
- ratfish/rabbitfish
shark properties
- fusiform streamlined body shape
- forward propulsion from trunk and caudal fin
- dorsal fin acts as stabiliser
- paired pectoral and pelvic fins for tilt
- no swim bladder
- hylostylic jaw extension, upper jaw is only attached by ligaments so sharks can grip and pull back prey
- cloaca
cloaca
single external opening for reproduction and waste removal
buoyancy in sharks
- have no swim bladder so buoyancy is maintained through combination of oil in their liver and continuous swimming
- continuous swimming also forces water across gills, keeping them oxygenated
- when sharks are resting on bottom of sea floor, they open and close jaws and use muscles in pharynx to squeeze and pump water across the gills
sensory physiology of sharks
- sharp black and white vision
- olfaction through nostrils (dead-end cups)
- electrophysiology for detecting electrical fields around them such as muscle contraction of prey
- no eardrums, entire body transmits sound to inner ear
Feeding mechanisms of sharks
- The largest sharks and rays are suspension feeders e.g. basking sharks
- Most are carniverous, so have several rows of teeth
- Teeth move to anterior as old ones are lost
- Have a shorter digestive tract than many vertebrates
- Evolved spiral valve to help with digestion of meat, a corkscrew-shaped ridge in intestine that increases surface area and slows digestive transit
reproductive strategies of sharks
- various reproductive strategies
- oviparous, ovoviviparous or viviparous
Oviparous
= eggs in protective coat hatch outside body
Ovoviviparous
= fertilised egg retained in oviduct, nourished by yolk, offspring hatch in uterus
Viviparous
= offspring develop in uterus, nourished by mother, birth live young
Rays
- Different lifestyle to sharks
- Mostly bottom dwellers, feed on molluscs and crustaceans
- Flattened shape
- Enlarged pectoral fins for propulsion
- Whiplike tail, many have venomous barbs
Osteichthyses
- Include bony fish and tetrapods
- almost all have an ossified endoskeleton with a hard calcium phosphate matrix
- 2 clades which evolved in the Devonian era, Actinoperygii and sarcopterygii
properties of bony fish
- operculum
- swim bladder
- most have bony scales, different to tooth-like shark scales
- mucus secreting glands reduce drag
- lateral line system
- diverse reproductive strategies
operculum of bony fish
- Protective bony flap covering gill chamber
- Movement of the operculum and contraction of the muscles surrounding the gill chamber draws water through the mouth and pharynx and out between the gills
buoyancy of bony fish
- maintained by the movement of gases between the blood and the swim bladder
- very efficient
- swim bladder evolved from early lungs in some lineages
Actinopterygii clade
- ‘ray-finned fish’
- Bony rays support fins, modified for manouvering and defence
- Originated in freshwater then spread to seas
- Many then returned to freshwater
- Some species are both freshwater and sea e.g. salmon
Sarcoptergyii
- ‘lobe-finned fish’
- Pectoral and pelvic fins have rod-shaped bones surrounded by a thick layer of muscle
- Evolved in brackish water such as coastal wetlands, used lobe fins to swim and ‘walk’ across substrate
3 extant lineages - Coelacanths (actinistia)
- Lungfish (dipnoi)
- Tetrapods
Coelacanths (actinistia)
- part of Sarcopterygii clade
- Secondarily derived cartilagenous skeletons
- 2 species, originally thought to have become extint ~65mya
Lungfish (dipnoi)
- part of the Sarcopterygii clade
- Evolved in ocean, now all 6 species are freshwater species in stagnant ponds and swamps
- Have lungs connected to pharynx, come to surface to gulp air
- Also have gills, main gas exchange organ
- Adapted to dry seasons by burrowing in mud
- closest living relatives to tetrapods
Evolution of tetrapods
~360mya
- Pectoral and pelvic fins evolved into limbs and feet in one lobe-fin lineage
- Supports weight on land and transmits muscle generated forces to ground to allow for locomotion
- Lineage also retained adaptations for aquatic life
Devonian and Carboniferous
- Greater diversity of tetrapods emerged, most still tied to water
Acanthostega, evolution of tetrapods
~365mya
- Fully formed legs, ankles and digits
- Bones supporting gills
- Tail-rays supporting fin
- Weak pectoral and pelvic girdles
- Possibly unable to support body, hypothesis that it slithered out of water occasionally
Other changes to the tetrapod body plan
- Head separated from body by neck
- Originally 1 vertebrae, can only move head up and down
- Evolution of second vertebrae allowed side to side movement of head, useful for predator detection
- Bones of pelvic girdle fused to backbone, forces from hind legs can be transferred to the whole body
- Extant tetrapods have no gill slits, embryonic pharyngeal gill slits give rise to other structures
class amphibia
3 orders
- urodela (tailed)
- anura (tailless)
- apoda (legless)
Urodela
- Tailed, e.g. newts, salamanders
- Some entirely aquatic, others on land as adults or throughout life
- Walk with a side-to-side bending body motion
- Trot gait, diagonally opposed limbs
- Paedomorphosis e.g. Mexican axolotl
Anura
- Tailless e.g. frogs
- More specialised for moving on land
- Adults have powerful hind legs and elastic muscles
- Tongue for prey capture and feeding
- Often have distasteful/poisonous toxins for predator evasion
- Colouration for defence, warning/mimicry, camouflage
Apoda
- Legless e.g. caecelians
- Secondarily derived
- Nearly blind
- Tropical
- most burrow in moist forest soil
- Some species are freshwater
Life stages of amphibians
- Anuran larval stage = tadpole (aquatic herbivore)
- Metamorphose into adults, legs, lungs, external eardrums develop, digestive system adapts to carnivory, gills (and lateral line system in may species) disappear
- Urodele and apodan larvae are carniverous and look like the adults
Habitats of amphibians
- Most live in damp habitats or burrow under vegetation
- Keeps skin moist for gas exchange
- Some terrestrial species lack lungs (breathe only through skin and through oral cavity
Other adaptations of tetrapods for terrestrial life
- a ribcage to ventilate lungs
- More efficient than throat based respiration
- Amphibians supplement breathing through the skin with breathing with the buccal cavity
- May have allowed amniotes to abandon respiration via their skin
- They then could evolve less permeable skin, giving them more ability to conserve water and therefore live on land
reproductive strategies of amphibians
- Most have external fertilisation
- Lay shelless eggs in water/a moist environment
- Won’t lose water through dessication
- Various types of parental care including mobile nurseries (back, mouth, stomach brooding)
- Some ovoviviparous, some viviparous
main characteristic shared by amniotes
- evolution of a terrestrially adapted egg = amniotic egg
- 4 extraembryonic membranes: yolk sac, allantois, amnion, chorion
- Most reptiles and some mammals also have a shell (leathery or calcified)
- Unclear evolutionary reasons, but may have allowed size and strength of egg to increase
properties of amniotic egg
- shell, protection against damage/ water loss, permeable for gas exchange
- Amnion and chorion (surrounding embryo) are vascularised, allowing for gas exchange
- allantois allows for excretion of waste and is also vascularised
- albumen for nutrition and protection
- yolk for nutrition
Amniote fossils
- Ancestor ~370mya
- Found in dry climates
- Herbivores (grinding teeth)
- Predators
- no fossils of amniotic eggs found
properties of reptilian clade
- Keratinous scales, protection against damage and dessication
- Most lay shelled eggs on land, therefore have internal fertilisation
- Many are viviparous
- Most are ectothermic, behavioural thermoregulation
- Birds are endothermic, metabolic thermoregulation
Earliest living reptiles
- Oldest fossils ~310mya
Parareptiles - Large, stocky, quadripedal
- Plated for defence
- Extinct by end of triassic ~200mya
Diapsids
- Diversified as parareptiles were dwindling (Triassic)
- Most obvious derived diapsid character = pair of holes each side of the skull behind eye socket (temporal fenestrae)
- 2 main lineages, Lepidosaurs and Archosaurs
extinct lepidosaurs
Marine reptiles e.g. giant mososaurs
extant lepidosaurs
- Tuataras, lizards and snakes
- Modified diapsid skull
- 2 extant lineages, Sphenodontians and Squamates
Sphenodontians
- Tuataras, 1 species
- Lizard-like reptiles
- Relatives ~220mya over many continents
- Now only 30 islands off New Zealand, not found on main island due to rodent introduction
- ~50cm
- Eat insects, small lizards, bird eggs, chicks
squamates
- lizards and snakes
- ~7900 species
Lizards
- Mostly small
- Jaragua lizard is 16mm long
- Komodo dragons ~3m (Indonesia)
- Hunts deer and stalks wounded prey, toxins in bite
snakes
- Legless (secondarily derived
- Some species show remnants of pelvic girdle and limb bones
movement of snakes
- Normal, lateral bending (Sidewinding, concertina)
- Rectilinear, grips ground with belly scales and pushes self forward, useful for prey ambush
Extinct archosaurs
- On land dinosaurs diversified into 2 main lineages
- Ornithischians
- Saurischians
Ornithischians
- Herbivores
- Many have anti-predator defences such as tail clubs and horned crests
Saurischians
- Long necked giant tree browsers
- Theropods = bipedal carnivores
e.g. Tyrannosauras rex, bird ancestors
Extant archosaurs
Crocodilians, 23 extant species
(Alligators, crocodiles and gharials)
evolution of crocodilians
- Emerged during Triassic
- Early crocodilians were small, terrestrial quadripeds with long slender legs
- Later adapted to aquatic habitats, evolution of upturned nostrils
properties of tesdudines
- Turtles, terrestrial and aquatic species
- Anapsid, no temporal fenestra
- Box-like shell
- Head retraction
Earliest turtles could not retract heads
Retraction mechanisms evolved independently
Pleurodires
Fold neck horizontally
Cryptodires
Fold neck vertically
testudine shells
- Most are hard for defence
- Clavicles and ribs fused to carapace (dorsal side of shell)
- Plastron = ventral side of shell
- Absence of transitional fossils to show evolution of shell
- Molecular data suggests that testudines are related to crocodilians
- Plates of archosaurs possibly became more extensive and started to form a shell
- Other data suggests they are perhaps a sister group to birds
Head retraction in testudines
- earliest turtles could not retract heads
- Retraction mechanisms evolved independently
- Pleurodires fold neck horizontally
- Cryptodires fold neck vertically
origin of birds
- Cladistic analysis from fossils shows they evolved from theropods
- Feathered theropod fossils show feathers evolved before flight
- Feathers possibly for insulation, mating or camouflage
Evolutionary theories of flight in theropods
- Cursorial (ground-up): small running dinosaurs gained lift from feathers, aided predatory lifestyle
- Arboreal (trees down): dinosaurs climbed trees and glided, aided by feathers
Theropods evolved into birds by 150mya
Archaeopteryx:
- Feathered wings
- Ancestral characteristics (teeth, clawed digits in wings, long tail)
Later Cretaceous bird fossils:
- Lost teeth and clawed forelimbs
- Acquired short feathered tail
extant birds
- neornithese clade
- 11000 species
- 32 orders
- variety of forms
- most have similar fusiform body shape adapted to flight
- beak shape highly adapted to diet
- foot structure highly adapted to lifestyle
neornithese clade, derived characteristics and modifications for flight
- wings with feathers made of beta keratin
- no urinary bladder
- females of most species have one ovary
- males have small gonads apart from during breeding season
- toothless (secondarily derived)
Advantages of flight
- Enhances hunting and scavenging
- Escape mechanism
- Ability to migrate long distances
birds, evolution of high metabolic rate
- as flight is energetically expensive
- Evolution of endothermy
- Feathers and fat for insulation
- Lungs have tubes to elastic air sacs, improving respiration
- 4 chambered heart
Order Sphenisciformes
- Flightless penguins
- powerful pectoral muscles
Ratites
- In order Struthinioformes
- Flightless, adaptations for cursorial locomotion include long legs
- Sternal keel absent (attachment for flight muscles)
- Small pectoral muscles
E.g. ostrich, emu, kiwi etc
synapsids
- evolutionary origin of mammals
- one lowered temporal fenestra on either side of skull
- 3 radiations of lineage
1st radiation of synapsid lineage
- Palaeozoic era
- evolution of pelycosoaurs (primitive)
- include sailbacks and sphenacodontids
- Beginning of 3 boned middle ear in sphenacodontids
2nd radiation - Paleozoic
- evolution of therapsids (more derived)
- Upright posture, limbs drawn in so they can be used like levers, also helps to free up respiratory muscles
- Increased jaw musculature
evolution of cynodonts
- Triassic
- Most derived therapsid, but still not a true mammal
- E.g. Triethelodon
- Size reduction to rat sized
- Evidence of endothermy
- turbinate (nasal) bones, increases airflow, reduces water loss
- Secondary palate, can swallow and breathe at the same time
- Reduced ribcage restricted to anterior allowed evolution of diaphragm
- Still conflict between hearing a chewing as jaw bone involved in channeling sound (mammalian inner ear bones evolved from synapsid jaw joint)
3rd radiation of synapsids
- Cenozoic
- Evolution of true mammals
- Earliest mammals very small, shrew sized
- Lived alongside dinosaurs until late Triassic
Features of early mammals
- Small, <100mm
- Nasolacrimal duct
- Evolution of fur for insulation
- Precise occlusion = molars fit together, helps with mastication
- Diphyodonte = two sets of teeth
- Not good at evaporative cooling
- Ears lack pinna (outer part)
- Low metabolism, suited to nocturnal lifestyle (cooler at night)
- Probably insectivorous and solitary
- Large olfactory lobes. more sophisticated sensory processing
- start of evolution of lactation
nasolacrimal duct
- Channel between tear ducts and nasal cavity
- Important for pheromones, olfactory sensing and cleaning eyes/fur
- characteristic anatomical feature of early mammals
early mammals, evolution of lactation
- No nipples, secreted on to fur
- Not yet a food source for young
- Antimicrobial, possibly for protecting eggs laid in nest
key features of mammals
- Hair/fur for insulation, communication, camouflage, sensation
- lactation and suckling
- variety of skin glands
mammalian skin glands
- Variety of skin glands
- Apocrine, eccrine and sebaceous
- Functions include lubrication, waterproofing, olfactory communication and thermoregulation
- Mammary glands evolved from skin glands
Suckling
- Unique mammalian feature
- reflex
- Must be able to make a seal with mouth and breathe and swallow at the same time
- Facial muscles required
- evolution of mobile lips and cheeks, used for other functions like communication later in evolutionary history
benefits of lactation and suckling
- Paternal care not required
- Offspring do not rely on seasonal food supply
- Provides care outside uterus (marsupials)
evolution of lactation and suckling alongside precise occlusion and diphyodonty
- lactation and suckling evolved before precise occlusion and diphyodonty
- Diphyodonty precedes precise occlusion
- Feeding on milk requires no teeth so reduces number of sets of teeth needed
- Having no teeth allows jaw growth and eruption of permanent teeth in an almost adult sized jaw
major mammalian lineages
- Allotheria (extinct multituberculates)
- prototheria
- theria
prototheria
- infraorder Ornithodelphia
- order monotremata
- oviparous (lay eggs)
- cloaca
- shoulder girdle has retained ventral elements, secondarily derived sprawling stance
Platypus
- prototherian
- 1 species
- Semi-aquatic
- Eat aquatic invertebrates
- Australian
- Venomous spurs
- Leathery ‘beak’ with electrosensitive pores
Echidna
- prototherian
- Insectiverous, long sticky tongue
- Short-nosed echidna
1 species
Australian
Eats ants and termites - Long-nosed echidna
3 species
New Guinea
Eats earthworms
Metatheria
- Traditionally order marsupiala, now 4 recognised orders
- Geographically split
- 1 order in New World (Americas)
- 3 orders in Australia
- Inflected jaw
- Epipubic bones retained
- Primitive dental formula: I5/4 C1/1 P3/3 M4/4
Australian methatherian orders
- Dasyurids, carnivores e.g. Tasmanian devil
- Peramelids, insectivorous e.g. bandicoots and bilbies
- Diprotodontians, modified lower incisors, 3 radiations: possums, koalas, kangaroos
theria
- metatheria and eutheria
- Viviparity = give birth to live young
- Nipples
- At least 2.5 coils of cochlea (improved hearing)
- Pinna = external ears
- Tribosphenic molars
- Urogenital and alimentary openings
- Derived shoulder girdle, ventral elements lost, scapula moves independently
- increases stride lengths, allows for specialised gaits
Eutheria
- May be referred to as placental mammals, but some metatheria have modified placentas
- Rapid radiation so phylogeny obscure
- Auditory bulla surrounds ear
- Herbivores had a post-orbital bar that stabilises the jaw
- Lost epipubic bones, possibly aiding the birth of larger offspring
- Primitive dental formula: I3/3 C1/1 P4/4 M3/3 (usually fewer incisors and molars than metatherians, but more premolars)
Biogeography of mammals
- Radiation peaked in cenozoic era as land masses separated
- vicariance and dispersal
Vicariance of mammals
- Populations separated by physical processes
- Distinct mammalian faunas on each continent
Dispersal of mammals
- Active movement of populations
- Mammalian faunas mixed when continents rejoined
Convergent evolution of therians
- Eutherian and metatherian mammals share body forms and habits
- Evolved similar characteristics to suit similar environmental conditions, no common shared ancestors
Humans, evolution of bipedal locomotion
- Frees up upper limbs
- Energy efficient running
- Ideal head position to scan horizon