“PROTOCHORDATES” & ORIGIN OF VERTEBRATES

Basic Chordate Characters: present at some stage in life cycle

Vertebrates are Chordates
Chordates are Deuterostomes
 Relationships of the Deuterostome Phyla
ECHINODERMATA HEMICHORDATA
ENTEROPNEUSTA PTEROBRANCHIA CHORDATA
(Acorn worms) (pterobranchs)

Burrowing lifestyles Sessile, colonial lifestyles

Pharyngeal pouches
Multiple pharyngeal slits Single pharyngeal slit
Notochord
Postanal tail
Endostyle or thyroid gland
Dorsal hollow nerve cord
Radial symmetry
Endoskeleton of calcite plates
Water vascular system
Ambulacral grooves with tube feet Tripartite body: proboscis
collar & trunk

Molecular & Larval

Similarities

Gill “skeleton”
Ciliated pharyngeal slits Novel Feeding Mechanism

Coelomates with “Deuterostome

Developmental Complex”
Basic larval similarities
 HEMICHORDATES
Tripartite body
Pterobranchs
Perforated pharynx* (sessile & colonial)

All suspension feeders Acorn worms

(solitary burrowers)

Not notochord homologue

“Acorn Worm” Burrowing, Detritus Feeder

Dorsal & ventral nerve cords

“Neurulated” collar
nerve cord in some
species
 Ciliary Suspension Feeding in Acorn Worms

Suspension vs. Filter

Feeders

Ciliary-Mucus
Feeding Mechanism
 Relationships of the Deuterostome Phyla
ECHINODERMATA HEMICHORDATA
ENTEROPNEUSTA PTEROBRANCNIA CHORDATA
(Acorn worms) (pterobranchs)

Burrowing lifestyles Sessile, colonial lifestyles Pharyngeal pouches

Multiple pharyngeal slits Single pharyngeal slit Notochord
Postanal tail
Radial symmetry
Endostyle or thyroid gland
Endoskeleton of calcite plates
Water vascular system
Dorsal hollow nerve cord
Ambulacral grooves with tube feet Tripartite body: proboscis
collar & trunk

Novel Locomotor Mechanism

Gill skeleton
Ciliated pharyngeal slits

Coelomates with “Deuterostome

Developmental Complex”
Basic larval similarities
 Relationships of Chordate Subphyla : Origin of Vertebrates

UROCHORDATES CEPHALOCHORDATES VERTEBRATES

Distinct head & brain (cephalization)

Greatly expanded pharynx Anteriorly extended notochord Muscularization of pharynx
Reduction of nervous system Oral cirri (tentacles) Special paired sensory organs (eyes, nose, ears)
Sessile Burrowing Neural crest tissue & neurogenetic placodes in embryo

Somites; segmented trunk muscles

Notochord & postanal tail retained in adult

BASIC CHORDATE FEATURES

 Adult Tunicate (Urochordate) Morphology
“Mouth”

Endostyle

Perforated
Endostyle Pharynx
Feeding Groove
Thyroid Homologue*

Flow Reversal
 Structure of Urochordate Larva

Photo detector
Gravity detector

Muscles not
segmented
myomeres
Urochordate Larval Metamorphosis

Residual Chordate Features

Perforated Pharynx

Endostyle
 Cephalochordate Adult Morphology

Segmented myomeres

Extended notochord

Oral ‘tentacles”
 Cephalochordates: Structure, Function, Lifestyle

Extended notochord Specialized Features Related to Burrowing

•  Anteriorly extended notochord
•  Muscularized notochord
• Atrium?

Atrium
Muscularized
notochord
Cephalochordates (live)
Urochordates (Live)
ORIGIN OF VERTEBRATES
 Relationships of Chordate Subphyla : Origin of Vertebrates

UROCHORDATES CEPHALOCHORDATES VERTEBRATES

Distinct head & brain (cephalization)

Greatly expanded pharynx Anteriorly extended notochord Muscularization of pharynx
Reduction of nervous system Oral cirri (tentacles) Special paired sensory organs (eyes, nose, ears)
Sessile Burrowing Neural crest tissue & neurogenetic placodes in embryo

Somites; segmented trunk muscles

Notochord & postanal tail retained in adult

BASIC CHORDATE FEATURES

 Relationships of Chordate Subphyla : Origin of Vertebrates

UROCHORDATES CEPHALOCHORDATES VERTEBRATES

Distinct head & brain (cephalization)

Greatly expanded pharynx Anteriorly extended notochord Muscularization of pharynx
Reduction of nervous system Oral cirri (tentacles) Special paired sensory organs (eyes, nose, ears)
Sessile Burrowing Neural crest tissue & neurogenetic placodes in embryo

Development of “New Head”

Somites; segmented trunk muscles

Notochord & postanal tail retained in adult

BASIC CHORDATE FEATURES

Feeding Strategies & Vertebrate Evolution

Large Prey Now Possible

Become Dominant Predators

Neural Crest Structures

Muscularized Pharynx &

Supportive Skeleton
Suction Feeding Begins

Ciliary Driven
Food Restricted to Zooplankton
 Neural Crest Tissue & Ectodermal (Neurogenic) Placodes

Key Innovation in Vertebrate History

See Text P. 186-187

 Developmental Innovation & the “New Head”

Ectodermal Placodes
ORIGIN OF VERTEBRATES
(And Fossil Record)
 Driving Forces (Selection) For Vertebrate Body Plan

Myxines (hagfish)

Increasing: Body Size & Aerobic Capacity (= sustained locomotion)

Cephalochordates
SURVEY OF VERTEBRATES: FISHES
Vertebrate Diversification & Time (Fossil Record)
Vertebrate Diversification & Time (Fossil Record)

Rise of Birds, Mammals

& Teleost fishes

Archosaurs Dominate

Rise of “Amphibians”

Primitive Fishes Dominate

 MYXINI

Cyclostomes
PETROMYZONTIFORMES
AGNATHA

CONODONTA

PTERASPIDOMORPHI

CEPHALASPIDOMORPHA

“ostracoderms” PLACODERMI
Relationships of Fishes (Classes)

CHONDRICHTHES

ACANTHODII
GNATHOSTOMATA

OSTEICHTHYES
Extant

Extinct
Class MYXINI (Hagfishes)

Primitive Features of modern Agnatha*

• No jaws
• No paired appendages
• No bone or mineralized tissues

(marine scavengers; carrion feeders)

* Plus other possible primitive traits in Mixini (see Handout)

Hagfish Movements & Feeding Tactics

Knotting behavior
 Class PETROMYZONTIFORMES

Lamprey (marine and fresh water predators / semi-parasites)

Endostyle
Larval stage: burrowing, suspension feeder
 Class PTERASPIDOMORPHI
Thoracic shield

Dermal Armor
All 3 basic vertebrate
hard tissues present
Extinct “pteraspids” • Enamel
• Dentine
• Bone (Aspidin)

Oldest “ostracoderms”
Class CEPHALASPIDOMORPHA = “Other Ostracoderms” in current text

Large Head Shields

Extinct
“Cephalaspids”

Fed on soft food

& detritus

Electro-receptors?

Shallow marine Pectoral lobes

habitats
 Class CEPHALASPIDOMORPHA

Anaspids: relationships to lampreys??

Reduced armor
Row lateral gill openings
Circular mouth

Probably Convergence!
Relationships of Jawed Vertebrates
 CLASS PLACODERMI Thoracic shields

First to have true jaws; Jaws enable

Vertebrates to become dominant predators Ambush predators?
 PLACODERMS
(Arthodires)

Major Predators of
Devonian Seas

Hinged between head &

Thoracic armor plates

Scale

Dunkleosteous
PLACODERMI
(Antiarchs)

First to successfully
invade continental waters

Might be first air breathing vertebrates

Arthrodire : Late Devonian of Australia

Hinge
Arthrodire : Late Devonian of Australia

Note: no ossified internal skeleton

Class CHONDRICTHYES: (modern) Subclasses: Elasmobranchii & Holocephali

No bone
External gill openings

Active marine predators

No swim or air bladders

 Pronounced
heterocercal
tail

Terminal Broad based fins

mouth

Enlarged pectoral
fins
 Specialized, deep water
shellfish feeders

Scale-less,
Open lateral line canals
 Holocephalian (“Ratfish”)

“Open”
Lateral line system
Internal Fertilization in Most Chondricthyians

“Claspers”on pelvic fins of males

 Elasmobranchs: large predators (marine mammal specialists)

Miocene
Great white

Modern Great white

 Class ACANTHODII
Prominent leading
edge spines

Covered gill openings

Small, marine
Late Devonian Seas Cladoselache & Acanthodians
Class OSTEICHTYHES : BONY FISHES
Class OSTEICHTHYES
Subclass ACTINOPTERYGII
Superorder PALAEONISCIFORMES
Superorder NEOPTERYGII Grades (not clades)

Teleosts: most numerous of

all vertebrates in modern world
 PALAEONISCIFORMES (“Chondrosteans”)

Typically: heavy scales

heterocercal tails
long gapes
poorly ossified skeletons
Sturgeon = chrondrostean (primitive actinopterygian)
 Chondrosteans : poorly ossified internal skeleton

Cartilage (blue)
Bone (red)
 Primitive NEOPTERYGII (“Holosteans”)
Relicts in Mississippi - Missouri Drainage

Reduced heterocercal tails

Well-ossified internal skeleton
Heavy scales
 Advanced Neopterygians: Teleosts

Salmon & trout:

primitive teleosts

Typically: homocercal tails

reduced scales
protrusible jaws
strongly ossified skeletons
 Advanced Neopterygians: Teleosts

Guppy

Enormous size range

Trenendous structural, behavioral
& ecological diversity

Marlin