Bio 370 Lecture 16: Chordates I

Phylogeny of Deuterostomia

Hollow dorsal
nerve cord

Fig. 14.1

Phylogeny of chordates

Key questions regarding evolution of chordates

1) Who were the first fishes?
2) Who were the direct ancestors of the first fishes?

= paraphyletic

Fig. 15.2

Phylum Chordata
The phylum includes vertebrates (animals with
backbones), but not all chordates are vertebrates

Cephalochordata

Urochordata

Vertebrata

Lancelets

Tunicates

Vertebrates

~ 29 species

~ 1600 species

~ 50,000 species

Phylum Chordata
Phylum named for the
notochord, a semirigid
rod of fluid-filled cells
enclosed in a fibrous sheath
Typically extends the length of body
ventral to the central nervous system
Supports & stiffens the body, provides
skeletal scaffold for swimming muscles
Present throughout life in some
groups, restricted to early
development in most
Fig. 15.1

Phylum Chordata: Homologies

All chordates have these
homologous structures:

(1)Dorsal nerve cord

(2) Notochord
(3) Pharyngeal slits
(4) Post-anal tail

In lancelets, these
structures are all
present in the adult

In most chordates,
some are present
only in the embryo

(5) Endostyle

Postanal tail

Fig. 15.7Adult lancelet

Representative chordate embryos

All chordates have these
homologous structures:

(1) Dorsal nerve cord

(2) Notochord
(3) Pharyngeal slits
(4) Post-anal tail
(5) Endostyle
In lancelets, these
structures are all
present in the adult

In most chordates,
some are present
only in the embryo

Phylum Chordata: Homologies

In vertebrates, the notochord, pharyngeal slits are present
only during embryonic development
The notochord is replaced by the backbone in adults

The pharyngeal slits develop into various structures:

- In fishes, they develop into parts of the jaws & gills
- In tetrapods, they give rise
to the Eustachian tube,
middle ear cavity, tonsils,
& parathyroid glands

Phylum Chordata: Homologies

The thyroid gland of vertebrates is homologous to the
endostyle of lancelets, tunicates, & larval lampreys

Many adult vertebrates have tails, but we lose ours!

Tail

Human embryo at 8 weeks

Human embryo at 10 weeks

Ancestral
chordate

Chordates

Lancelets

Tunicates

Head

Vertebral column

Jawed vertebrates

Sharks,
rays

Ray-finned
fishes

Jaws

Lobe-fins
Lobe-finned
fishes

Lungs or lung derivatives

Lobed fins

Reptiles

Legs
Amniotic egg

Mammals
Milk

Amniotes

Simplified
phylogeny of
chordates

Tetrapods

Amphibians

Vertebrates

Lampreys

Evolutionary time

Neural crest

Craniates

Hagfishes

Phylum Chordata
Subphylum Urochordata
Tunicates: sea squirts & others
About 1600 species

Live in all seas, at all depths

As adults, most are sessile,
baglike, suspension feeders

The adult body is enclosed in

a tough, nonliving tunic made
of proteins & cellulose
From 1 mm to 1 m long

Phylum Chordata
Subphylum Urochordata
Incurrent siphon: anterior
Excurrent siphon: dorsal
Pharynx fills most of body
Anus empties into the
atrium inside the tunic
Heart & some blood
vessels present
Hermaphroditic
Gametes exit via the excurrent
siphon, fertilization is external

Phylum Chordata
Subphylum Urochordata
Water enters the pharynx via
the incurrent siphon, passes
through the pharyngeal slits
into the atrium, & exits via
the excurrent siphon
Endostyle: a ciliated groove
on the ventral side of pharynx
- Secretes a sheet of mucus
that traps food particles
- Mucus drawn into esophagus
Fig. 15.4

Phylum Chordata
Subphylum Urochordata
Tunicate larvae have all five chordate homologies
Larva swims for several hours before attaching to an object &
metamorphosing into a sessile adult
During metamorphosis, the notochord & tail disappear, & the
dorsal nerve cord is reduced to a ganglion above the pharynx

Fig. 15.5

Phylum Chordata
Subphylum Cephalochordata
Lancelets
Genus Branchiostoma,
formerly Amphioxus
About 29 species worldwide

Live in sandy bottoms of

coastal waters
From 3-7 cm long

Phylum Chordata
Head
Notochord
Mouth

Dorsal,
hollow
nerve cord

Pharynx
Endostyle
Pharyngeal slits

Subphylum Cephalochordata
All five chordate homologies
are present in adult lancelets
Like vertebrates:
- Muscles in V-shaped segments
- Fishlike circulatory pattern
Unlike vertebrates:
- No brain, cranium, vertebral
column; few sense organs

Digestive tract
Atriopore

Segmental
muscles
Anus
Post-anal tail

Phylum Chordata
Subphylum Cephalochordata
Cilia in the buccal cavity & pharynx draw water in through the
mouth
Water passes through the pharyngeal slits, where food particles
are trapped in mucus secreted by the endostyle
Mucus & food are moved by cilia into the gut

Fig. 15.7

Phylum Chordata
Subphylum Cephalochordata
As in tunicates, water enters
the atrium via the pharyngeal
slits, exits atrium through
the atriopore
Gas exchange occurs over
the whole body surface (not just
in the gills)
Separate sexes
Gametes exit via the atriopore,
fertilization is external

Cross-section of a lancelet

Phylum Chordata
Subphylum Vertebrata
Vertebrates: animals with vertebral columns
All vertebrates are also craniates:
they have a skull enclosing the brain
In our textbook, Vertebrata = Craniata
But hagfishes (Myxini), which have a
cranium, do NOT have vertebrae
In cladistic classifications, hagfishes are
considered craniates but not vertebrates:
Craniata = Myxini + Vertebrata

Adaptations in Vertebrate Evolution

Earliest vertebrates were larger & more active than lancelets &
tunicates (protochordates)
Modifications of skeletal structures & muscles permitted increased
speed & mobility
The higher activity level & larger size of vertebrates required
adaptations for:
- Locating, capturing, & digesting food
- Supporting a high metabolic rate

Vertebrates
Musculoskeletal Modifications
The endoskeleton permits almost unlimited body size

The skeleton probably was composed initially of cartilage

- All vertebrate embryos have skeletons made of cartilage
- Hagfishes, lampreys, & sharks have cartilaginous skeletons

Vertebrates
Musculoskeletal Modifications

Segmented body muscles (myomeres or myotomes) changed

from V-shaped muscles of cephalochordates to W-shaped
muscles of vertebrates

More complex folding of vertebrate myomeres provides powerful

control over a long body

Vertebrates
Musculoskeletal Modifications
The extracellular matrix of bone tissue is
made of collagen & hydroxyapatite, a
mineral that contains calcium & phosphate
ions
Bones original function may have been
mineral storage
Phosphorus & calcium are used in many
physiological processes

In high demand in organisms with high

metabolic rates

Vertebrates
Musculoskeletal Modifications
Structural strength of bone is superior to cartilage ideal for muscle
attachment in areas of mechanical stress
Early fishes such as ostracoderms featured a body covered with
plates of bone
Many ray-finned fishes have bony scales
Bony fish scales are typically thin

Fig. 15.4

Vertebrates
Physiological Modifications
Digestive, respiratory, circulatory, excretory systems are
modified to meet increased metabolic demand
Early chordates used pharyngeal slits for filter-feeding

In vertebrates, the addition

of muscles to the pharynx
created a powerful pump
With the origin of highly
vascularized gills, gas
exchange became the main
function of the pharynx

Vertebrates
Physiological Modifications
A ventral, chambered heart & erythrocytes with
hemoglobin made transport & exchange of
gases & other substances more efficient
To manage the increased
amounts of food ingested:
- Muscular action replaced
ciliary action for moving
food through the gut
- Accessory digestive glands
(liver & pancreas) produced
secretions to aid digestion

Vertebrates
Nervous System Modifications
Shifting from sessile filter feeding to active predation
required new sensory & motor, & integrative controls
Anterior end of nerve cord enlarged as a tripartite brain
(Forebrain, midbrain, & hindbrain)
Brain protected by a cranium
of cartilage or bone
Paired sense organs
for vision, equilibrium,
& sound evolved

Vertebrates: Neural Crest & Ectodermal Placodes

The head & sense organs develop from two embryonic
structures found only in vertebrates:
Ectodermal placodes: plate-like
thickenings of the ectoderm on
either side of the neural tube
Give rise to the lens of the eye, olfactory
& inner ear epithelia, & other structures
Neural crest: bilateral bands of
ectodermal cells lying along the
embryonic neural tube

Vertebrates: Neural Crest & Ectodermal Placodes

The notochord forms
from mesoderm along
the embryos axis
Notochord later replaced
by the vertebral column
The neural plate forms
from ectoderm above
the notochord

Neural
fold

Notochord
Ectoderm
Mesoderm
Endoderm

Neural folds

Neural
plate

Vertebrates: Neural Crest & Ectodermal Placodes

The neural plate rolls up to form
the neural tube, which sinks
below the embryos surface
The underlying notochord induces
the formation of the neural tube
The neural tube will become the
brain & spinal cord

Neural
fold

Neural plate

Outer layer
of ectoderm

Neural tube

Vertebrates: Neural Crest & Ectodermal Placodes

Dorsal edges
of neural plate

Neural
crest

Ectoderm

Neural
tube
Ectoderm

Neural crest cells

migrate to distant
sites in the embryo
The cells give rise to some of
the bones & cartilage of the
skull & face, portions of the
eyes, ears, & teeth, parts of the
nervous & endocrine systems,
& other structures

Notochord

Migrating neural
crest cells

Chordate Evolution & the Position of Amphioxus

Chordate evolution has taken two paths:
1) Sedentary urochordates
2) Active, mobile cephalochordates & vertebrates
In 1928, Walter Garstang suggested that the ancestral chordate
was a sedentary filter-feeder, like an adult tunicate
- The swimming larva failed to
metamorphose into a sessile adult
- Instead, it developed gonads & began
reproducing in the larval stage

Chordate Evolution & the Position of Amphioxus

Garstang called this process paedomorphosis: the evolutionary
retention of juvenile traits in the adult

But phylogenetic studies & fossils

place cephalochordates as sister to
urochordates + vertebrates
Also, neural crest tissue was
recently found in urochordates

Fig. 15.10

Chordate Evolution & the Position of Amphioxus

Thus Garstangs hypothesis has been falsified:
- The ancestral chordate was probably a free-swimming
filter-feeder like amphioxus
- The sessile body form of urochordates is unique
But amphioxus lacks features that were present in the most recent
common ancestor of vertebrates:
- No tripartite brain, chambered
heart, special sensory organs,
muscular gut & pharynx,
or neural crest tissue

Ancestral
chordate

Chordates

Lancelets

Tunicates

Neural crest

Head

Vertebral column

Jawed vertebrates

Sharks,
rays

Ray-finned
fishes

Jaws

Lobe-fins

Lungs or lung derivatives

Lobed fins
Reptiles

Legs
Amniotic egg

Mammals
Milk

Amniotes

Simplified phylogeny
of chordates

Tetrapods

Amphibians

Vertebrates

Lampreys

Craniates

Hagfishes

Phylogeny of chordates

= paraphyletic

Fig. 15.2

Ancestral Chordates
Pikaia , a middle
Cambrian fossil from
the Burgess Shale
(~530 Mya) is one of
the earliest known
chordates

Pikaia had a notochord

& V-shaped myomeres

Fig. 15.8

Ancestral Chordates
Haikouella, recently discovered at Haikou, China
Chordate features: notochord, pharynx, & dorsal nerve cord
Vertebrate features: pharyngeal muscles, paired eyes,
& an enlarged brain
But it lacks a cranium, so it cannot be a vertebrate

Fig. 15.9

Ancestral Chordates
Conodonts are a group of
fossil aganthans
Once known only from small
fossils of their bony teeth
Recent fossils suggest
conodonts were eel-like fishes,
but maybe not vertebrates
Condonts may have possessed
a pair of large eyes, tapering
tail, a simple fins with fin rays,
chevron-shaped muscles, & a
notochord

Top left: conodont elements on head of a nail. Top

right: conodont elements. Note how these elements
seem to form a bilateral set. Bottom: an artists
depiction of a conodont. The condodont specimens
known to date range from ca. 1-40 cm long.

Early Vertebrates
Ostracoderms are paraphyletic assemblage of bony fishes
Plates of bone in their skin and lacked jaws
Some with paired fins, some without paired fins
Sucked water into pharynx by muscular pumping
Some species were dorso-ventrally flattened while others were cylindrical

Fig. 15.11

Late Cambrian (500 mya) to the end of Devonian (359 mya)

Evolution of Jaws from Cartilaginous Gill Arches

Fossil & embryological evidence suggests that jaws arose from
the 1st or 2nd cartilaginous gill arches

Upper jaw

In this Carboniferous
shark, the jaws resemble
the gill supports

A similar transformation
occurs during
development
in modern sharks
Fig. 15.13

Lower jaw

Gnathostomes: jawed vertebrates (living & extinct)

Spiracles (one on each side of the

head of some fishes) are remnants of
the second gill openings (openings
between mandibular and
hyomandibular arches)

spiracle

Spiracles are possessed by some

species of sharks, but especially rays

internal gill opening

Early Jawed Vertebrates

Placoderms were among the first jawed vertebrates (Gnathostomes)
- Armored with diamond-shaped scales or plates of bone
- Late Silurian (420 mya) to the end of Devonian (359 mya)

Fig. 15.14

Acanthodians with multiple paired fins

Extinct group with characteristics in common with sharks and bony
fishes

Evolution of Paired Pectoral & Pelvic Appendages

All gnathostomes also have paired pectoral & pelvic
appendages in the form of fins or limbs

Early Jawed Vertebrates

Some species of placoderms were quite
large
Some species of Dunkleosteus grew to a
length of 6 m and featured a large mouth
with chisel-shaped teeth
Interestingly, some placoderm species are
only known from their cranial armor and
jaws
Given the rounded head with teeth
adapted for sheering, species
Dunkleosteus were probably open-water
(pelagic) predators

Dunkleosteus sp. (Arthrodira): the massive

head and anterior trunk plates (together
almost the size of a small car) of
Dunkleosteus sp., a placoderm